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Annals of Tropical Medicine 
and Parasitology 




THE UNIVERSITY OF LIVERPOOL 


Annals 

OF 

Tropical Medicine and 
Parasitology 


ISSUED BY THE 

Liverpool School of Tropical Medicine 

Edited by 

Professor J. W. W. STEPHENS, M.D.Cantab., D.P.H. 

Professor R. NEWSTEAD, M.Sc., J.P., F.R.S., A.L.S., F.E.S., Hon. F.R.H.S. 

_yorke, m.d. 

Annals of Tropical Medicine and Parasitology 

Vol. VIII, p. 585, lines 8-10. The authors, Drs. Breinl and 
Young, wish to amend ‘Any .... dryness;’ to read thus:—‘Any 
lead sulphide was filtered off, washed with sulphuretted hydrogen 
‘water, dissolved in dilute nitric acid, and the excess of acid 
‘ removed by repeatedly evaporating to dryness 

With Frontispiece , thirty-nine plates , twenty-pve figures in text, menty-jour 

charts, and three maps 


LIVERPOOL: 

AT THE UNIVERSITY PRESS, 57 ASHTON STREET 


Editorial Secretary 
Dr. H. B. FANTHAM, 
School of Tropical Medicine , 
The University , 
Liverpool . 



CONTENTS 


No. i. March 18, 1915 

Bayon, H. page 

Leprosy : A Perspective of the Results of Experimental Study of the Disease. 

Plates I-VI . 1 

Young, W. J. 

The Metabolism of White Races living in the Tropics. I.—The Protein 

Metabolism . 91 

Ward, Henry B. ; and ITirscii, Edwin F. 

The Species of Paragonimus and their Differentiation. Plates YII-XI ... 109 

ScmvETZ, Dr. J. 

Preliminary Notes on the Mosquitos of Kabinda (Lomami), Belgian Congo... 163 
Stephens, J. W. W. 

On the Peculiar Morphological Appearances of a Malaria Parasite. Plate XII 169 
Carter, Henry F. 

On Some Previously Undcscribed Tabanidac from Africa. Plate XIII . . 173 

Seidelin, Harald. 

Experiments with Salvarsan-Copper in Trypanosomiasis . 197 

Stephens, J. W. W. ; and Stott, W. 

Studies in Blackwater Fever. III.—The Relationship of Quinine to Blackwatcr 

Fever . 201 

CONTENTS 

No. 2. June 30, 1915 

Brf.inl, Anton. pace 

Gangosa in New Guinea and its Etiology. Plates XIV-XVII . 213 

Priestley, Henry. 

TheiUria tachyglossi (n.sp.) a Blood Parasite of Tachyglossus aculcatus. Plate 

XVIII. 233 

Scott, H. Harold. 

An Investigation into the Causes of the Prevalence of Enteric Fever in Kingston, 

Jamaica ; with Special Reference to the Question of Unrecognised Carriers 239 



CONTENTS No. 2 — continued 


Rrkinl, An ion. page 

On the Occurrence and Prevalence of Diseases in British New Guinea. Plates 

XIX-XXVI . 285 

Fantham, H. B. 

Inject Flagellates and the Evolution of Disease, with Remarks on the Importance 

of Comparative Methods in the Study of Protozoology ... ... ... 335 

CONTENTS 

No. 3. July 31, 1915 

PACK 

Yorke, Warrington ; and Blacklock, B. 

Notes on the Bionomics of Glossina palpalis in Sierra Leone, with Special 

Reference to its Pupal Habitats. Plates XXVII-XXXIII . 349 

Yorke, Warrington; and Blacklock, B. 

Food of Glossina palpalis in the Cape Lighthouse Peninsula, Sierra Leone ... 363 

Fraser, Henry ; and Fletcher, W. 

The Cultivation of the Leprosy Bacillus. 381 

Yorke, Warrington ; and Blacklock, B. 

The Reservoir of the Human Trypanosome in Sierra Leone. 383 

Fantham, H. B. 

Spirocbaeta hronchialis , Castcllani, 1907, together with Remarks on the Spiro- 

chaetes of the Human Mouth. Plate XXXIV . . 391 

Yorke, Warrington ; and Blacklock, B. 

Notes on Certain Animal Parasites of Domestic Stock in Sierra I,cone ... 413 

Davey, J. B. 

The Etiology of Juxta-articular Subcutaneous Nodules ... ... ... 421 

Yorke, Warrington ; and Blacklock, B. 

Ankylostomiasis in Dogs in Sierra Leone. 425 

Stephens, J. W. W. 

Studies in Blackwatcr Fever. IV.—Note on a Case of Quartan Malaria 

associated with Blackwater Fever . 429 



CONTENTS 


No. 4. December 30, 1915 

PAGE 

Macfie, J. W. Scott 

Nuclear Variations of the Ncutrophilc Leucocytes (Arnetli Counts) in Malaria 


and Yellow Fever . 435 

Macfie, J. VV. Scott 

Babesiasis and Trypanosomiasis at Accra, Gold Coast, West Africa. Plates 

XXXV, XXXVI . 457 

Breinl, A. ; and Priestley, H. 

Differential Counts and the Neutrophile Blood-Picture of Natives-—Adults 

and Children—of New Guinea . 495 

Macfie, J. W. Scott 


A Case of Dysentery in a Monkey, in which Amoebae and Spirochaetcs were 

found. Plate XXXVII . 507 

Schwetz, Dr. J. 

Preliminary Note on the General Distribution of Glossina pal pahs , Rob-Dcsv., 


in the District of Lomami, Belgian Congo. 513 

Macfie, J. W. Scott 

A Note on a Trypanosome of the Black Rat (Epimys rattus). Plate XXXVIII 527 
Bayon, H. 

The Artificial Cultivation of Hansen’s ‘ Bacillus’ . 535 

Stephens, J. W. W. 

Studies in Blackwater Fever. V.—The Duration of Haemoglobinuria ... 539 

Fantham, H. B.; and Porter, Annie 

Some Experimental Researches on Induced Herpetomoniasis in Birds. 

Plate XXXIX. 5+3 

Lloyd, Ll. 

On the Association of VVarthog and the Nkufu Tick ( Ornithodorus moubata) ... 559 

Porter, Annie 

On Anaplasma-like Bodies in the Blood of Vertebrates ... 


561 









Volume IX 


March, 1915 


No. 1 


ANNALS 

# OF 

TROPICAL MEDICINE AND 
PARASITOLOGY 


ISSUED BY 

THE LIVERPOOL SCHOOL OF TROPICAL MEDICINE 


Edited by 

Professor J. W. W. STEPHENS, M.D. Cantab., D.P.H. 

Professor R. NEWSTEAD, M.Sc., J.P., F.R.S., A.L.S., F.E.S., Hon. F.R.H.S. 
Professor WARRINGTON YORKE, M.D. 

AND 

Professor Sir RONALD ROSS, K.C.B., F.R.S., M.D., F.R.C.S., 

Major I.M.S. (Ret.) 


Editorial Secretary 
Dr. H. B. FANTHAM, 
School of Tropical Medicine , 
The University , 
Liverpool . 



C. Tinting < 5 r* Co., Ltd. 

Printers to the University Press of Liverpool 
53 Victoria Street 



THE INCORPORATED 

LIVERPOOL SCHOOL OF TROPICAL MEDICINE 

Founded by Sir ALFRED LEWIS JONES, K.C.M.G. 

{Affiliated with the University of Liverpool) 

Hon. President: H.R.H. Princess Christian of Schleswig-Holstein. 

Chairman : MR. F. C. DANSON. 

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The Earl of Derby, K.G. 

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Sir William H. Lever, Bart. 

Mr. O. Harrison Williams 


COMMITTEE 


Mr. H. J. Read, C.M.G. 
Vice-Chancellor Sir Alfred Dale 
Sir W. B. Bowring, Bart. 
Professor R. CATON 
Professor W. HERDMAN, F.R.S. 
Professor J. M. Beattie 
Dr. C. J. Macalister 
M r. G. P. Newbolt 
M r. C. Booth (Jun.) 

Mr. T. F. Harrison 
Mr. A. R. Marshall 
Mr. W. Roberts 
Mr. J. W. Alsop 
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Mr. C. Livingston 


Colonial Office 
University of Liverpool 

■ Council of University of Liverpool 
| Senate of University of Liverpool 
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Mr. A. L. Rea, Hon. Treasurer 

Mr. A. H. MILNE, C.M.G., Secretary. 

B io Excharge Buildings, Liverpool 



Staff, 1915 

1. At the University of Liverpool 

Professors - - JOHN WILLIAM WATSON STEPHENS, M.D., 

Cantab., D.P.H. Sir Alfred Jones Professor of 
Tropical Medicine 

ROBERT NEW STEAD, E.R.S., M.Sc., A.L.S., 
F.E.S., Dutton Memorial Professot of Entomology' 
LiF.rT.-Coi., Sir RONALD ROSS. K.C.B., F.R.S', - 
LL.D., E.R.C.S., D.P.H., M.D., D.Sc. Nobel 
Laureate 1902. (Indian Medical Service, retired>. 
Professor of Tropical Sanitation 
WARRINGTON YORKE, M.D.. If alter Myers 
Professor of Parasitology 

Lecturers - Prof. E. W. HOPE, M.D., D.Sc., Lecturer or 

Municipal Sanitation 

HENRY FRANCIS CARTER, S.E.A.C. Dim.. 

F.E.S., Lecturer on Entomology 
HAROLD BENJAMIN FANTHAM, M.A., D.Sc., 
Lecturer on Parasitology 

WILLIAM THOMAS PROUT, M.B , C.M.G., 
Lecturer on Tropical Sanitation 
HAROLD WOLFERSTAN THOMAS, M.D., 
C.M. 

Honorary Lecturer - Major JOSEPH FITZGERALD BLOOD, M.D.. 

M.Ch. (Indian Medical Service, retired) 

Honorary Statistician WALTER STOTT 

2. At the Research Laboratories 

Director - - - B. BLACKLOCK, M.D., D.P.H. 

3. At the Royal Infirmary, Liverpool 

Physician - - JOHN WILLIAM WATSON STEPHENS, M I)., 

Assistant to the Cantab., D.P.H. 

Physician - WILLIAM THOMAS PROUT, M.B , C.M.G. 


4. At the Royal Southern Hospital, Liverpool 


Phys icians 


Surgeon 


- CHARLES JOHN MAC A LISTER, M.D., 

F.R.C.P. 

JOHN LLOYD ROBERTS. M.D.. M.R.C.P. 

- GEORGE PALMERSTON NEWBOLT, M B., 

F.R.C.S. 


Hon. Tropical 

Pathologist 


JOHN WILLIAM WATSON STEPHENS, M.D.. 
Cantab., D.P.H. 


5. At the Yellow Fever Research Laboratory of the School, 
Manaos 

Director - - HAROLD WOLFERSTAN THOMAS, M.D., 

C.M. 


Laboratory 

JOHNSTON LABORATORY, UNIVERSITY OF LIVERPOOL 


Hospitals: 

ROYAL INFIRMARY AND ROYAL SOUTHERN HOSPITAL, 
LIVERPOOL 


Secretary's Office: 

B 10, EXCHANGE BUILDINGS, LIVERPOOL 



NOTICE 


The following courses of instruction will be given by the 
Liverpool School of Tropical Medicine during 1916 :— 

Full Course begins 6 January. Advanced Course begins 1 June. 
Diploma Examination, 3 April. Certificate Examination, 30 June. 

Full Course begins 15 September. 

Diploma Examination, 11 December. 

These dates are subject to revision. 

The full Course of Instruction is open to all qualified medical men, 
and the Examination to all students who have taken out this full 
course. 

Fee for the full Course of Instruction—Thirteen Guineas. 

Fee for the Diploma Examination—Five Guineas. 

Fee for the Short Course of Instruction—Four Guineas. 

Fee for the use of a School microscope during one term—Ten 
shillings and sixpence. 

For prospectus and further information, application should be made 
to the Dean of the Medical Faculty, University of Liverpool. 


The following have obtained the Diploma in Tropical Medicine of 
the University of Liverpool: — 

Diploma in Tropical Medicine 


Date 0 / 

Difloma 

1904 Augustine, Henry Joshua 
1904 Bennett, Arthur King 
1904 Bruce, William James 
1904 Byrne, John Scott 
1904 Clayton, Thomas Morrison 
1904 Dalziel, John McEwen 
1904 Dee, Peter 

1904 Greenidge, Oliver Campbell 
1904 Hehir, Patrick 
1904 Khan, Saiduzzafor 
1904 Laurie, Robert 
1904 Maclurkin, Alfred Robert 
1904 McConnell, Robert Ernest 
1904 Nicholson, James Edward 
1904 Philipson, Nicholas 
1904 Sharman, Eric Harding 
1904 Thomson, Frank Wyville 

1904 Walker, George Francis Clegg 

1905 Anderson, Catherine F.lmslie 
1905 Brown, Alexander 

1905 Caldwell, Thomas Cathcart 
1905 Critien, Attilio 
1905 Hooton, Alfred 
1905 Hudson, Charles Tilson 
1905 Illington, Edmund Moritz 


Date oj 
Diploma 

1905 Macfarlane, Robert Maxwell 
1905 Maddock, Edward Cecil Gordon 
1905 Moore, James Jackson 
1905 Nightingale, Samuel Shore 
1905 Radcliffe, Percy Alexander Hurst 

1905 Young, John Cameron . 

1906 Adie, Joseph Rosamond 
1906 Arnold, Frank Arthur 
1906 Bate, John Brabant 
1906 Bennetts, Harold Graves 
1906 Carter, Robert Markham 
1906 Chisholm, James Alexander 
1906 Clements, Robert William 
1906 Dundas, James 

1906 Faichnie, Norman 
1906 Jeffreys, Herbert Castelman 
1906 Mackenzie, Donald Francis 
1906 Pailthorpe, Mary Elizabeth 
1906 Palmer, Harold Thornbury 
1906 Pearse, Albert 
1906 Sampey, Alexander William 
1906 Smithson, Arthur Ernest 
iqo6 Taylor, Joseph van Someron 
1906 Taylor, William Irwin 
1906 Tynan, Edward Joseph 



Date of 
Difloma 

1906 Watson, Cecil Francis 
1906 Willcocks, Roger Durant 

1906 Williamson, George Alexander 

1907 Allan, Alexander Smith 
1907 Allwood, James Aldred 
1907 Bond, Ashton 

1907 Branch, Stanley 

1907 Collinson, Walter Julius 

1907 Davey, John Bernard 

1907 Donaldson, Anson Scott 

1907 Fell, Matthew Ilenry Gregson 

1907 Gann, Thomas William Francis 

1907 Graham, James Drummond 

1907 Hiscock, Robert Carroll 

1907 Keane, Joseph Gerald 

1907 Kennan, Richard Henry 

1907 Kenrick, William Hamilton 

1907 Le Fanu, George Ernest Hugh 

1907 Mackey, Charles 

1907 Maddox, Ralph Henry 

1907 McCarthy, John McDonald 

1907 Raikes, Cuthbert Taunton 

1907 Ryan, Joseph Charles 

1907 Vallance, Hugh 

1908 Caverhill, Austin Mack 
1908 Crawford, Gilbert Stewart 
1908 Dalai, Kaikhusroo Rustomji 
1908 Dansey-Browning, George 
1908 Davidson, James 

1908 Dickson, John Rhodes 
1908 Dowdall, Arthur Melville 
1908 Glover, Henry Joseph 
1908 Greaves, Francis Wood 
1908 Goodbody, Cecil Maurice 
1908 Harrison, James Herbert Hugh 
1908 Joshi, Lemuel Lucas 
1908 Le Fanu, Cecil Vivian 
1908 Luethgen, Carl Wilhelm Ludwig 
1908 Mama, Jainshed Byramji 
1908 McCay, Frederick William 
1908 McLellan, Samuel Wilson 
1908 Pearce, Charles Ross 
1908 Schoorel, Alexander Frederik 
1908 Smith, John Macgregor 
1908 Stewart, George Edward 

1908 Tate, Gerald William 
1008 Whyte, Robert 

1909 Abercrombie, Rudolph George 
1909 Allin, John Richard Percy 
1909 Armstrong, Edward Randolph 
1909 Barrow, Harold Percy Waller 
1909 Beatty, Guy 

1909 Carr-White, Percy 
1909 Chevallier, Claude Lionel 
1909 Clark, William Scott 
1909 Cope, Ricardo 
1909 Fleming, William 
1909 Hanschell, Hother McCormick 
1909 Hayward, William Davey 
1909 Henry, Sydney Alexander 
1909 Innes, Francis Alexander 


Date of 
Diploma 

1909 Jackson, Arthur Frame 
1909 Kaka, Sorabji Manekji 
1909 McCabe-Dalfas, Alfred Alexander 
Donald 

1909 Meldrum, William Percy 
1909 Murphy, John Cullinan 
1909 Samuel. Mysore Gnananandaraju 
1909 Shroff, Kawasjee Bvramjee 
1909 Thornely, Michael Harris 
1909 Turkhud, Violet Ackroyd 
1909 Webb, William Spinks 
1909 Yen, Fu-Chun 


1910 Brabazon, Edward 

1910 Castellino, Louis 

1910 Caulcrick, James Akilade 

1910 Dowden, Richard 

1910 Haigh, William Edwin 

1910 Hamilton, Henry Fleming 

1910 Hefferman, William St. Michael 

1910 Hip well, Abraham 

1910 Homer, Jonathan 

1910 Houston, William Mitchell 

1910 James, William Robert Wallace 

1910 Johnstone, David Patrick 

1910 Korke, Vishnu Tatyaji 

1910 Macdonald, Angus Graham 

1910 Macfie, John Wm, Scott 

1910 Manuk, Mack Walter 

1910 Murison, Cecil Charles 

1910 Nanavati, Kishavlal Balabhai 

1910 Nauss, Ralph Welty 

1910 Oakley, Philip Douglas 

1910 Pratt, Ishmael Charles 

1910 Sabastian, Thiruchelvam 

1910 Shaw, Hugh Thomas 

1910 Sieger, Edward Louis 

1910 Sousa, Pascal John de 

1910 Souza, Antonio Bernardo de 

1910 Waterhouse, John Howard 

1910 White, Maurice Forbes 

1911 Blacklock, Breadalbane 

1911 Brown, Frederick Forrest 

1911 Chand, Diwan Jai 

1911 Holmes, John Morgan 

1911 levers, Charles Langley 

1911 lies, Charles Cochrane 

1911 Ingram, Alexander 

1911 Kirkwood, Thomas 

1911 Knowles, Benjamin 

1911 Liddle, George Marcus Berkeley 

1911 Lomas, Emanuel Kenworthy 

1911 Mackarell, William Wright 

1911 MacKnight, Dundas Simpson 

1911 Mascarenhas, Joseph Victor 

1911 Murray, Ronald Roderick 

1911 Oluwole, Akidiva Ladapo 

1911 Rao, Koka Ahobala 

1911 Sinton, John Alexander 

19 n Tarapurvalla, Byramji Shavakshah 

1911 Taylor, John Archibald 

1911 Woods, William Medlicott 




Date of 
Diploma 

1912 Aeria, Joseph Reginald 
1912 Anderson, Edmund Litchfield 
1912 Borle, James 
1912 Bowie, John Tait 
1912 Brassey, Laurence Percival 
19 1 2 Christie, David 
1912 Dillon, Henry de Courcy 
1912 Dunn, Lillie Eleanor 
1912 Hardwicke, Charles 
1912 Jagose, Jamshed Rustomji 
1912 Kochhar, Mela Ram 
1912 McGusty, Victor William Tighe 
1912 Milne, Arthur James 
1912 Mitra, Manmatha Nath 
1912 Myles, Charles Duncan 
1912 Pelly, Huntly Nevins 
1912 Prasad, Bindeshwari 
1912 Prentice, George 
1912 Ross, Frank 
1912 Russell, Alexander James 
Hutchison 

1912 Ruthven, Morton Wood 
1912 Sandilands, John 
1912 Seddon, Harold 
1912 Smalley, James 
1912 Strickland, Percy Charles 
Hutchison 

1912 Watson, William Russel 

1913 Austin, Charles Miller 
1913 Banker, Shiavux Sorabji 
1913 Becker, Johann Gerhard us 
1913 Carrasco, Milton 

1913 Clark, James McKillican 

1913 Forsyth, Charles 

1913 Grahame, Malcolm Claude Russell 

1913 Grieve, Kelburne King 

1913 Hargreaves, Alfred Ridley 

1913 Hepper, Evelyn Charles 

1913 Hiranand, Pandit 

1913 Jackson, Oswald Egbert 

1913 IChaw, Ignatius Oo Kek 

1913 MacKelvie, Maxwell 

1913 MacKinnon, John MacPhail 

1913 Macmillan, Robert James Alan 


Date of 
Diplotna 

1913 Mouat-Biggs, Charles Edward 
Forbes 

1913 Noronha, John Carmel 
1913 O’Connor, Edward 
1913 Olubomi-Beckley, Emanuel 
1913 Pestonji, Ardeshir Behramshah 
1913 Puttanna, Dodballapur Sivappa 
1913 Reford, John Hope 
1913 Smith, Edward Arthur 
1913 Stewart, Samuel Dudley 
1913 Walker, Frederick Dearden 
1913 Wilbe, Ernest Edward 
1913 Wilson, Hubert Francis 
1913 Yin, Ulg Ba 

1913 Young, William Alexander 

1914 Arculli, Hassan el 

1914 Chohan,Noormahomed Kasembha 
1914 Connell, Harry Bertram 
1914 Gerrard, Herbert Shaw 
1914 Gimi, Hirji Dorabji 
1914 Gwynne, Joseph Robert 
1914 Hodkinsou, Samuel Paterson 
1914 Jackson, Arthur Ivan 
1914 Kaushash, Ram Chander 
1914 Kelsall, Charles 
1914 Luanco y Cuenca, Maximino 
1914 Misbah, Abdul-Ghani Naguib 
1914 Naidu, Bangalore Pasupulati 
Balakrislma 

1914 Rowe, John Joseph Stephen 
1914 Roy, Raghu Nath 
1914 Sbiveshwarkar, Ramchandra 
Vishnu 

1914 Sur, Sachindra Nath 
1914 Talati, Dadabhai Cursedji 
1914 Wilkinson, Arthur Geden 

1914 Wright, Ernest Jenner 

1915 Lobo, John Francis 
1915 Madbok, Gopal Dass 
1915 Pearson, George Howorth 
1915 Swami, Karumuri Virabhadra 
1915 Wood, John 





ANNALS OF TROPICAL MEDICINE 
AND PARASITOLOGY 


EDITORIAL NOTICE 

Articles for publication are accepted, if found suitable, from any 
source and will be understood to be offered alone to these Annals. 
They should be typewritten and addressed to: —The Editorial 
Secretary, School of Tropical Medicine, The University, Liverpool. 

Illustrations for text figures should be drawn clearly and firmly 
in Indian ink, if possible, on Bristol board. 

All lettering, names or legends on text-figures, charts or maps 
should be printed sufficiently large to allow of clear legibility on 
reduction if necessary. 

Plates and illustrations should be accompanied by short explana¬ 
tions. 

References to authors in the text must be made in the following 
way:—'According to Smith (1900) the spleen is enlarged, but 
Robinson (1914) says the reverse.’ The references should be 
collected in alphabetical order of authors’ surnames at the end of 
the paper, and arranged in the following way : — 

Robinson. S. (1914). ‘The spleen in malaria.' Annah of Nosology, 
Vol. XX. pp. 20-25. 

Smith, J. (1900). ‘ Enlargement of the spleen in malaria.’ Journal of 

Pathometry , Vol. I, pp. 1-20. 

Fifty reprints are supplied to each author, free of charge. 
Additional copies (up to 200) can be supplied at cost price. 


Subscription: £\ 2s. 6d. per volume, post free, payable in 
advance to the Secretary, Liverpool School of Tropical Medicine, 
B 10 Exchange Buildings, Liverpool. Correspondence concerning 
advertisements should also be addressed to the Secretary. 



LEPROSY: A PERSPECTIVE OF THE 
RESULTS OF EXPERIMENTAL STUDY 
OF THE DISEASE 


BY 

H. BAYON, M.D., 

RESEARCH BACTERIOLOGIST (LEPROSY), GOVERNMENT OF THE UNION OF SOUTH AFRICA, 

AT ROBBEN ISLAND 


(Received for publication 18 August , 1914) 


Plates I—VI 

CONTENTS 


I—Introduction. 

II —The Clinical Nosology or Leprosy . 

III— Microscopical Features or Leprous Lesions . 

IV — Morphology and Staining Properties or Hansen's 4 Bacillus * 

Analogy with the Mycobacterium tuberculosis . 

V —Artificial Cultivation or the Micro-organism or Leprosy 

VI— Experimental Transmission to Animals. 

Rat-Leprosy and its Relation to the Human Disease . 

VII —Serological Tests and Reactions in Leprosy. 

VIII —Communicability and Contagion. 

Is the transmission through an intermediate arthropod probable ? 

IX— Early and Difperential Diagnosis. Prognosis . 

X— Treatment, Specific and Palliative . 

XI —Prevention. Home and Asylum Segregation. Settlements 

XII —Discussion or Results and Conclusions. 

Explanation or Plates . 


4 

9 

18 

20 

23 

3 1 

40 

47 

55 

6o 

64 

70 

73 

77 

8 o 


I. INTRODUCTION 

Leprosy is the name applied to all clinical manifestations 
produced by an infection with the micro-organism commonly known 
as ‘Hansen’s bacillus’ {Mycobacterium leprae). 

It is a disease that has been recognised and feared for centuries, 
possibly already as far back as 2400 B.C., or in any case about 
1300 B.C. (Ebers papyrus). Leprosy as described in Leviticus does 





2 


not, however, correspond exactly to the symptoms we know in more 
modern times. It should, moreover, not be forgotten that the word 
' leprosy ’ is but an approximate rendering of the expression 
‘zaraath’ in the original Hebrew. 

Not only is leprosy the ‘ most ancient disease,’ but it is also a 
widely spread, cosmopolitan scourge, which nearly rivals con¬ 
sumption, syphilis and cancer, if not in numbers, at least in the 
variety of climatic conditions under which it can flourish and 
increase. Though at the present day it is found, to any con¬ 
siderable extent, practically only in tropical and sub-tropical 
countries, it is known that numerous lepers existed in France, 
Germany, Great Britain, and other countries of Europe throughout 
the Middle Ages. Opinions differ whether it was imported into 
Europe from the East through the returning Crusaders. The 
disease still lingers in Turkey, Russia, the Balkan States, 
Norway, Spain, Portugal, parts of Italy, etc., in the form of small 
endemic foci. However, the ruthless individual and, in some cases, 
collective segregation in the thirteenth and fourteenth centuries, 
aided no doubt by the improving hygienic conditions of all classes, 
succeeded in bringing about a practically complete extinction of 
leprosy in Middle Europe and Great Britain. In addition to this, 
various plague epidemics also helped in sweeping to an early grave, 
in a somewhat selective fashion, the vagrant and pauper lepers 
sooner and quicker than individuals belonging to better situated 
classes. 

The slow if intermittent course of the disease, the extensive 
mutilations and repulsive features, which are an outstanding 
symptom of its more advanced stages, have rendered leprosy a 
subject of terror and horror from time immemorial. That it is 
contagious has been known for centuries, but it is no doubt mainly 
due to aesthetic reasons that the fear it engendered was quite out 
of proportion to the degree in which it is communicable, under 
usual circumstances, from the diseased to the healthy. 

Leprosy presents numerous peculiarities in its pathognomonic 
features. Not only is its infectivity extremely variable under slightly 
dissimilar or even identical circumstances, but its period of incuba¬ 
tion varies also within very wide limits, anything between one year 
and twenty-five or thirty. The ‘causa prima morbi’ has been 



3 


known for years and years, and yet the fundamental bases of all 
bacteriological research, viz., artificial cultivation of the micro¬ 
organism and the experimental transmission of the disease, have 
presented insurmountable difficulties up to the present day. 
Indeed, these bases are still matters under discussion, and in any 
case successful results can only be achieved after numerous, 
repeated, prolonged, pertinacious attempts. The large numbers of 
bacteria which may be present in single leprotic lesions are certainly 
not surpassed in any other disease, with the possible exception of 
bubonic plague. 

An excellent instance showing the influence that bacteriology 
exercises on clinical medicine is to be found in the effects resulting 
from the discovery of the ‘ bacillus ’ of leprosy by Armauer Hansen 
in 1872. It is true that definite knowledge of the morphology of 
the micro-organism was only obtained after Neisser had succeeded 
in demonstrating it by means of an ‘acid-fast' stain in 1879. 
However, from the moment the bacterial origin of the disease was 
recognised, the communicability of leprosy became scientifically 
established, and all doubts in that connection could only be the 
outcome of an incomplete or lacking comprehension of the applica¬ 
tion of the laws of biology to practical medicine. 

On the other hand the extreme difficulty encountered in 
transmitting the disease to animals, the repeated lack of success 
attending the inoculation of leprous material into human beings 
(with one exception: Aming), have allowed much doubt to arise 
regarding the exact fashion in which leprosy is transmitted. Even 
the purely bacteriological observation of the fact that apparently 
the ‘bacilli’ could not be artificially cultivated, and isolated from 
the lesions they produced, has seriously hampered the therapeutic 
outlook. 

Leprosy slowly and surely became the disease of negations; it 
could not be experimentally transmitted, its bacterium could not be 
isolated in artificial culture, and so to complete the list, leprosy 
could not be treated, let alone arrested, and no hope of cure could 
be entertained. 

It is my intention to show whether recent experimental work 
allows, enables, or justifies us in taking a more active and positive 
view of the situation. 



II. THE CLINICAL NOSOLOGY OF LEPROSY 

Two clinical types of the disease are generally distinguished 
and no more. Such views are partly owing to the influence of 
Hansen, who considered that it was not necessary to make further 
sub-divisions because practically all lepers presented features at one 
time or another of their existence (that is, if they lived long 
enough) which allowed them to be classed either as cases of 
(i) nodular (hypertrophic) leprosy, or (ii) maculo-anaesthetic, other¬ 
wise called atrophic, smooth leprosy. 

From a purely clinical point of view this classification holds 
good, however, only within certain limits. It is a commonly 
accepted rule of nosology that bedside or clinical nomenclature 
does not take into consideration so much the eventual course of a 
disease, as its etiology and the symptoms and appearances which, 
however fleeting and changeable, can be brought under a definite 
scheme of relative incidental constancy. The classification used in 
relation to recognised clinical types of tuberculosis, syphilis, and 
other complaints, which often run a chronic course and vary largely 
in their symptomatology, seems clearly to show which are the 
determining factors ruling the designation of clinical entities. 

It is no doubt quite correct, from a general point of view, to 
state, as Hansen did, that a ‘mixed’ leper (i.e., a patient who from 
the very beginning shows features of both the nodular and maculo- 
anaesthetic type) who lives long enough and undergoes a natural 
process of elimination of the great majority of the acid-fast 
‘ bacilli ’ from his lesions, will eventually be classifiable as a purely 
anaesthetic leper. Many of these instances of * lepra mixta,’ 
however, do not reach this ultimate stage, because death steps in 
before completion of the process of evolution into a different type, 
and in any case the primary condition may last for many years. 

Without unnecessarily labouring this point any further, I 
believe that there is quite sufficient justification in considering every 
single case of leprosy as classifiable on the basis of the clinical and 
bacterioscopic features at the time of examination, irrespective of 
the expected course of the disease later on. In other words, the 
clinical diagnosis is to be considered of a temporary nature, as far 
as the naked eye appearances (and to a certain extent also the 
bacterioscopic results) are concerned. The diagnosis leprosy is the 



5 


only unchangeable feature, though it may need modification by the 
adjectives ‘ arrested ’ or possibly even ‘ cured.’ 

Leaving aside sub-divisions founded on partial or purely 
localised symptoms, I believe that on examination of any great 
number of lepers, in any part of the world, more or less numerous 
specimens of the following varieties or even distinct types of the 
disease may be distinguished. 

1. Nodular type with more or less marked or diffuse tuberosities 
which generally are most developed on the face, but of course can 
also affect any other part of the body, including (though somewhat 
rarely) the scalp, the soles of the feet, the tongue, and the palms 
of the hands. Leprous nodules should contain numerous acid-fast 
micro-organisms, and in certain instances may be located as loose 
or movable small lumps which can be shelled out from under the 
skin. 

Nodular leprosy may give rise to visceral lesions, which are 
remarkable for the enormous quantity of acid-fast ‘ bacilli ’ that 
can be detected accumulated in the tissues of the spleen, liver, 
testicles, lymphatic glands and lungs. 

2. Maculo-anaesthetic type with more or less numerous spots 
which have a slight predilection for the trunk (chest, back, buttocks) 
but are certainly also very frequently present on the limbs. The 
spots in advanced cases may show a greater or lesser amount of 
anaesthesia. It should be noted that often the anaesthesia is 
definitely localized in portions not showing any maculae. In many 
cases, however, the anaesthesia is by no means complete, and should 
more correctly be termed paraesthesia. The patients cannot 
distinguish hot and cold, sharp or blunt in the affected areas, but 
are still sensitive to touch. In early stages the maculae may present 
a ringed appearance; the centre then may be normal in relation to 
sense of touch, etc., whilst the outer ring, if anything, is slightly 
hyperaesthetic. 

The maculae appear to be the result of the localisation (and 
possibly disintegration) of a small number of casual micro¬ 
organisms in the skin. The anaesthetic areas are due to the 
‘ bacilli ’ invading the sheath of the nerves, causing a proliferation 
of connective tissue, which brings about a slow degeneration of the 
corresponding nerve fibres. In advanced cases the bacilli invade 



6 


the ganglia of the spinal cord, hence the practical impossibility, 
under certain circumstances, of distinguishing leprosy clinically 
from syringomyelia. 

5. Mixed or combined leprosy, that is, cases where from the 
very earliest noticeable onset of the disease, and for years after¬ 
wards, definite areas of paraesthesia (insensibility to heat and cold, 
incapacity of distinguishing blunt from sharp) and anaesthesia can 
be made out, accompanied by maculae and small nodules teeming 
with acid-fast bacteria. 

An excellent description of this type has been given by Gluck, 
who called it ‘ lepra tubero-anaesthetica.’ 

These three main types can be sub-divided into the following 
relatively rare varieties and singular forms: 

1. Lupoid or serpiginous leprosy. Among the 2,000 lepers I 
have had under observation in South Africa I have only met with 
the disease three times. Another instance of the variety was seen 
in London. It is recognisable as leprosy by the presence of nerve- 
lesions, but acid-fast micro-organisms are very scarce indeed in any 
of the superficial patches. The expression ‘ patches ’ is used 
because the skin-lesions are much more raised and thickened than 
mere ‘ maculae,’ and yet do not deserve the appellation of nodules, 
being too flat and extensive. They spread by forming peculiar 
confluent configurations, which can be best recognised by referring 
to PI. V, fig. 1. 

The patches are a peculiar coppery colour, and do not show any 
marked tendency to break down and ulcerate, which in addition to 
other peculiarities distinguishes them from lupus. 

At times single patches whose colour, texture and general 
appearance corresponds closely to that of these serpiginous lesions, 
are noticeable in cases of leprosy, whose remaining clinical features 
differ in no other fashion from the usual nodular type. 

In other instances the patches may not be confluent and 
serpiginous, but are distributed unevenly all over the body, varying 
in size from the diameter of a threepenny-bit to the extent of a 
man’s hand or more. They may appear relatively rapidly, within 
a fortnight or so, and are accompanied by localised paraesthesia. 
In such case the diagnosis usually presents considerable difficulty, 
because acid-fast bacteria may be practically absent in the lesions 
and nasal mucosa. 



7 


Jadassohn has described somewhat similar varieties of leprosy 
as the ‘ tuberkulid ’ type, in view of the resemblance which micro¬ 
scopical specimens of the lesions bear to tuberculosis of the skin. 

2. Many cases of maculo-anaesthetic leprosy begin with 
paraesthesia, numbness, and a peculiar ‘ leathery ‘ feeling, usually 
of the ring and little finger of one hand, or tingling sensations with 
small areas of definite anaesthesia may be present in a hand or 
foot. Such conditions can last for a considerable time, as much as 
three or more years, before definite maculae make their appearance. 
Some cases, after a prolonged premonitory stage of this nature, may 
develop nodules and finish as definite hypertropic leprosy. 

Maculo-anaesthetic lepers may heal in such a fashion as to allow 
hardly any traces of their spots being visible to the naked eye; the 
trained observer may just be able to make out a simple configurated 
discoloration or slight livido. Needless to say, definite areas of 
anaesthesia are usually easily made out in these patients. As long 
as these conditions last it appears quite correct to classify them as 
simple paraesthetic or anaesthetic leprosy. 

3. Under certain circumstances maculae or spots may appear 
before any nerve symptoms are traceable. Even to the experienced 
clinician the leprotic nature of the spots may appear doubtful, 
other conclusive symptoms being absent and possibly no history of 
contact being available, though the latter has but a relative value. 
In these rare occasions only prolonged observation, aided and 
controlled by the microscope, can afford us certainty in diagnosis. 
Eventually, in the course of time, a definite maculo-anaesthetic 
stage is reached, but during this initial period the condition is one 
of simple macular leprosy. 

When dealing with dark-skinned races great care must be taken 
not to mistake for macular leprosy one of the numerous tineae which 
have been so ably classified and investigated by Castellani. 

4. In some instances leprotic symptoms are seen which are 
somewhat too prominent to deserve still the name of ‘ maculae ’ or 
spots. The edges are considerably more raised than is usually the 
case in smooth leprous lesions, owing to active proliferation of the 
superficial dermal tissue. The periphery is studded with numerous 
minute papulae, which have a somewhat vesicular appearance. The 
central portion may or may not participate in this process of 
granulation, but a certain degree of desquamation is generally 



8 


present. In any ease the appearance of the inner area is angrier and 
more ‘ active-looking ’ than what is seen in the usual run of maculae, 
though parts of the surface do not usually participate to such a 
marked degree in the process of granulation and eruption notice¬ 
able at the margin. 

It may be objected that practically all maculae of lepers go 
through periods of increased activity or exacerbation, during which 
they may show some of the appearances described. However, the 
case illustrated in PI. V, fig. 2, has remained in this condition 
for over three years, and moreover, the localisation of the lesion 
is somewhat peculiar. In addition, the maculae of many lepers 
never reach the condition of marginate proliferation shown in the 
photograph. 

Of course this whole condition is but the result of an intensified 
tissue reaction, such as generally takes place to a much lesser 
degree in most maculae. The diagnosis, however, may still 
present difficulties, because, as in the instance illustrated, other 
definite leprotic symptoms, such as anaesthesia, may be absent or 
cannot be detected at the time of examination. I should like to 
draw attention to the existence of these intensified ‘maculae* 
without insisting on their separate classification. 

5. Diffuse or hyperaemic leprosy is also scarcely a separate 
variety, but simply an initial or premonitory stage of nodular 
leprosy. It consists in a peculiar diffuse, puffy swelling of the 
eyebrows, the lobes of the ears and the alae nasi, which in addition 
takes on a peculiar pinkish-bluish hue. This condition may last for 
years and imperceptibly merge into the well-known condition of 
leprosy generally known as ‘ facies leonina.’ In rare instances the 
disease may not make further progress, but eventually improves and 
leaves, as the only trace, a peculiar wrinkled and dry appearance of 
the skin of the face. I have so far not found any of such arrested 
cases without smaller or larger areas of anaesthesia. 

These three types and five varieties or peculiar diversities from 
the usual clinical appearances of leprosy appear to me to cover all 
forms of leprosy, as far as I have been able to observe them. In 
addition, there are numerous combinations which can result through 
atypical and transitory appearances of the disease mixing in varying 
proportions. 



9 


Bullae, ulcers, suppurations and necrosis may considerably alter 
and add complexity to the clinical features of the symptomatology 
of leprosy. They should, however, be considered as complications 
of the primary infection; moreover, they are too variable and 
inconstant to allow definite systematization. 

Careful description of the majority of these symptoms is also 
given by Abraham (1910) where special stress is laid on the fact, 
and rightly too, that the diagnosis is in many cases but the 
description of a stage of the disease. 

These various sub-divisions have not been made to complicate 
further a diagnosis which under usual circumstances is already 
sufficiently difficult, but on the contrary to induce a careful observa¬ 
tion of the various phenomena of a disease whose correct, exact, and 
above all early recognition is at all times a matter of considerable 
importance. 

On two occasions I have noticed in lepers peculiar psoriasis-like 
maculae, but in these instances the possibility of psoriasis and 
leprosy occurring together could not be excluded. 

REFERENCES 

Abraham, Ph. (1910). Leproiy in * System of Medicine,* edited by Allbutt-Rolleston, Vol. II, 
Part II, pp. 648-694. 

Balfour, A. (1913). Leprosy in 4 Tropical Diseases Handbook, Ghent Exhibition,* pp. 90-104. 

Castellani and Chalmers (1913). Leprosy in 4 Manual of Tropical Medicine,’ 2nd edition, 
pp. 1147-1169. 

Jadassohn, J. (1913). Lepra in 4 Handbuch der pat. Mikro-organismen,* pp. 791-930. 

Manson (1911). Tropical Diseases, pp. 527-565. 


III. MICROSCOPICAL FEATURES OF LEPROUS LESIONS 

The minute anatomy of leprotic lesions or histo-pathology of 
leprosy is the study, by means of the microscope, of the relationship 
or direct and indirect action of the lepra-bacterium to and on the 
tissues it has invaded. Accordingly, every investigation or analysis 
of the appearances detected by the microscope must take into 
consideration not only the presence or absence of acid-fast ‘ bacilli,’ 
but also, in the former eventuality, the quantity and localisation of 
the ‘ bacilli ’ discernible in the sections being examined. Accordingly 
it is to be understood that the microscopical diagnosis of a leprous 
lesion is bacterioscopic and histo-analytic. 



10 


Good descriptions of the microscopical appearances of leprosy 
have been given by Lie, MacLeod, Jadassohn and others, and an 
even prolonged observation of the material available in South 
Africa does not bring to light any new facts of fundamental 
importance, except to show once more the necessity of clearly 
distinguishing between leprosy and tuberculosis by means of 
animal experiments and any other methods available for the 
purpose. In referring to the lesions found in the spleens of lepers, 
MacLeod (1910) expresses his view as follows (page 314): 

'In the spleen, in addition to leprous infiltrations in the 
trabeculae, pulp and malpighian corpuscles, extensive necrotic 
masses have been described by Arning (Schaffer, Lepra, 1900, I, 
p. n) like those of tuberculosis with typical Langhans giant cells. 
Such cases have to be further verified from the point of view of 
tuberculosis before they can be finally accepted/ 

In several autopsies (5) at Robben Island I have found in the 
spleen of nodular lepers lesions similar to, if not identical with, those 
described and illustrated by Schaffer. The inoculation experiments 
into guinea-pigs were, however, positive; that is, the animals 
experimented upon died within two to four months with extensive 
generalized lesions of a tuberculous nature. 

These investigations are by no means concluded, and they have 
the limited value of single instances. They simply show that 
‘ tuberculous 9 lesions in a leper are not necessarily always caused by 
Hansen's ' bacillus 9 alone. 

The uncompromising standpoint of Hansen, that leprosy and 
tuberculosis were so absolutely distinct from each other that no 
single common feature was thinkable, is no more defensible or 
convenient, not even from a didactic point of view. At the present 
day we have learnt through the more intimate knowledge of many 
maladies and the properties of numerous pathogenic micro¬ 
organisms, that dissimilar diseases can be caused by microbes which 
are practically indistinguishable, and that on the other hand 
analogous pathological conditions can be brought about by micro¬ 
organisms which behave in a totally different fashion in artificial 
culture and laboratory tests. As cases in point it is only necessary 
to mention at random yaws and syphilis; kala-azar, infantile 
splenomegaly and oriental sore; typhoid and paratyphoid; human 



II 


and bovine tuberculosis, plague and pseudo-tuberculosis. Pathology 
has learnt the truth of Goethe’s dictum ‘ Alles ist Ubergang.’ The 
increasing exactitude of clinical methods of observation ought to 
enable us to avoid the pitfall of considering two allied diseases as 
identical, since we understand that hard and fast types are only 
extant for the purpose of classification. 

For some time discussion has been rife whether leprotic tissue 
ever contained giant cells of the Langhans type. It is now practi¬ 
cally unanimously admitted by those who have had opportunity of 
seeing a sufficient number of slides from different cases, that giant 
cells do occur in typical leprotic tissues, for they have been 
observed by Thoma, Jadassohn, Ramon y Cajal, Rikli, Schaeffer, 
Dohi, Babes, Kedrowsky, Matsuda, Gurd, Bayon and G. W. 
Robertson (verbal communication). Even Lie, who originally held 
that giant cells were only present in leprosy when the disease was 
complicated by tuberculosis, now admits their presence, with the 
reservation that they are not morphologically absolutely identical 
to the Langhans type found in tuberculosis (verbal communication). 

In discussing the nosology of leprosy, stress was laid on the 
temporary nature of many conditions in leprosy, as a result Of the 
extremely chronic course of the disease and the facility with which 
it can be complicated by other infections. In addition we have to 
consider the multifarious appearances of the disease per se. These 
factors influence also the histological features to a considerable 
extent, so that it is not possible to state, except in a somewhat 
general fashion, which are the microscopical features that render 
the diagnosis ‘ leprosy ’ certain in any section. No single element is 
of decisive value, except the presence of numerous acid-fast rods, 
and this even with several qualifications. Hansen’s ‘bacillus* is 
generally only to be found in large numbers in advanced nodules 
and nerve-lesions, and is practically absent from maculae and 
leprotic papulae, that is, only to be detected after a very prolonged 
search and by special methods. 

Unna’s definition of a leproma is as follows: — 

‘ .... a diffuse granuloma, whose peculiarity consists, on 
the one hand, in its limitation to the connective tissue elements, and 
specially to the lymphatic system of the skin, and on the other in 
the enormous growth of organisms, whose number far exceeds 



\ / 


12 


anything we are accustomed to find in other infectious diseases. . . 
To the paucity of the cellular elements and the preponderance of 
the organisms dormant in the bacillary mucus, the remarkable 
indolence and relative benignancy of these growths may be 
ascribed.* (Unna-Walker; The Histopathology of the Diseases of 
the Skin, 1896, p. 616.) 

Jadassohn sums up his investigations on the subject in the 
following words* (l.c. p. 871): — 

‘The leprosy bacillus produces a non-characteristic type of 
inflammation, attended by the formation of granulomatous tissues 
with different features of a degenerative character, which can even 
deserve to be classified as necrosis; this may in some instances take 
on a tuberculous appearance, signalised by sclerosis, and direct or 
indirect destruction (Untergang) of various specific parenchymata. 
As a rule the leprosy bacillus causes only an indolent reaction and 
the resistance displayed by tissue against its luxuriant growth 
(Wucherung) and toxines is quite remarkable. Deposits of bacilli 
may be detected even in tissue which otherwise appears to be quite 
normal, and in cells which cannot be shown to be altered in any 
known fashion. The histological features are also very variable as 
a result of the diversity of clinical types.’ 

‘ The typical leproma, wherever it may be situated in the body, 
has the broad characteristics of other infective granulomata, with 
certain peculiarities which render its diagnosis possible ’ (MacLeod). 

It is evident from what has been said above that distinctive 
features need not necessarily be present in many lesions which from 
their occurrence and localisation we know to be definitely leprotic. 
By avoiding generalisation, and considering single types and 
anatomical situations by themselves, it is, however, possible to 
recognise certain definite, recurring, important features. 

Skin. 'Nodules. Numerous acid-fast rods occur, especially in 
matted groups, there is absence of advanced necrosis (except in the 
presence of open sores), and no caseation or giant cells with a 
necrotic centre. Giant cells containing single acid-fast ‘ bacilli ’ may, 
however, be present, but generally they are not numerous, and show 
fewer peripheral nuclei than is the case in tuberculosis. The 
‘ bacilli ’ may be scattered loosely in the parenchyma of the nodule 

* Translated by the Author of this paper. 



or may have invaded various cells of endothelial and fibroblastic 
origin, in which case it is especially remarkable that the structure 
of the protoplasm and nucleus or even the morphology of the 
bacterial acid-fast rods has not suffered in any noticeable fashion. 
Plasma cells are rare as opposed to yaws and syphilis, where they 
are especially numerous. 

A peculiar narrow band of hyaline, glassy tissue, containing no 
‘ bacilli,’ is often noticeable just under the epidermis. 

Acid-fast bacteria may be present in such quantities that in 
stained sections (Ziehl-Nielsen) the whole tissue appears a dull red 
to the naked eye. In other instances ‘ bacilli,’ though very 
numerous, are distributed singly or by twos and threes, and can 
only be detected with high-power lenses. 

Skin. Patches. Acid-fast bacteria are not numerous. They 
can only be detected with great difficulty. They are not bunched 
or massed, but present in little groups of few ‘ bacilli,’ which may 
apparently be situated extra-cellularly or they may have invaded 
cells without causing any alteration in the structure of either 
protoplasm or nucleus. 

Giant cells with numerous peripheral nuclei arc often present. 
At times a peculiar coagulation or discrete necrosis of the tissue 
may be noted in parts, which shows hardly any definite structure. 
It has a filamentary appearance and stains easily with eosin. Such 
small areas of degeneration are circumscribed and show one or two 
giant-cells at the periphery. 

The vessels and capillaries can be seen partaking in a process of 
proliferation, accompanied by an increase in the number of 
endothelial cells forming their walls. 

The epidermis is often found . thinner than usual, and the 
epithelial villi are somewhat flattened out, or the epithelium may 
appear as a border of uniform breadth. 

Spindle-shaped cells (young connective tissue cells) may be 
numerous, and plasma cells are not infrequent. 

Skin. Maculae. Nodules and patches partake of the nature of 
infectious granulomata. The macula belongs, however, to another 
pathological group, which can be compared with a condition of 
slight chronic irritation, possibly not very dissimilar to that seen in 
Erythema perstans. 



4 


Various degrees of perivascular cellular deposit correspond to 
the different intensity of eruption in single maculae, but, as a rule, 
a slight infiltration of spindle and round cells can be detected 
surrounding the capillaries and sweat-glands. The hair-follicles 
may also undergo a similar process of localised infiltration and 
proliferation. Giant cells may be present, but hardly any acid-fast 
bacilli, unless nodules are to be found in other parts of the body. 

The histological appearances I have described agree with the 
view that maculae are a local anaphylactic symptom, caused by the 
toxic products set free by the lepra-bacilli, after disintegration 
induced through the agency of the defensive properties of the 
tissues involved. This process takes place, however, when only few 
germs are present, because lepra ‘ bacilli ’ in common with other 
acid-fast micro-organisms are very resistant. 

Regarding the changes occurring in the peripheral nervous 
system, the suggestion has been made by Lie that ‘ bacilli ’ are only 
present in maculae at the very beginning, and that later they 
disappear into the nerve-endings and finally ascend along the 
peripheral nerves. It can. however, be objected that in many cases 
anaesthesia appears to set in without the primary production of 
maculae in the corresponding area, and that many maculae do not 
show any definite anaesthetic symptoms, though they have lasted 
for years. 

The other hypothesis considers the neuritis to be a descending 
process (a contrast to the ‘ ascending ’ process put forward by Lie), 
and accordingly the result of deposit following generalised 
infection. In any case, though the nerves may be involved in any 
single type of leprosy, it is in the maculo-anacsthetic type that this 
invasion and localisation gives rise to the most striking and 
apparently relatively rapid results. 

In thickened peripheral nerves, under certain circumstances, 
numerous ‘ bacilli ’ may be found, as in skin lesions. Generally 
they are situated in massed groups in the spindle-cells of the 
perineurium, or loose in the interstitial lymphatic spaces; at times 
they may be distributed with a certain regularity in the cells of the 
endoneurium, so that they appear to be in rows rather than in 
clumps. In the course of time they may get eliminated and leave 
behind a marked thickening, consisting of connective tissue with a 



5 


few stray degenerated and fragmented nerve fibres, in which the 
general disposition of a nerve-bundle is still recognisable. 

Practically every organ of the body has been found to be the 
site of leprotic lesions by some observer or other, but deposits of 
lepra-bacilli are more usually found in the lungs, liver, spleen and 
testes. In the lungs the frequent complication with tuberculosis is 
very disconcerting, and in many instances a decision as to the 
pathological process involved can only be arrived at by tests with 
inoculated guinea-pigs, though it is quite possible that even this 
crucial experiment may not have an absolutely decisive value. 
Babes gave considerable attention to the question of the lesions 
found in the lungs of lepers, and has come to the following 
conclusions which well deserve to be quoted in full : 

(1) The lungs are normal, and do not contain lepra bacilli. 

(2) They appear healthy to the naked eye and under the microscope, but 

notwithstanding contain leprosy micro-organisms. 

(3) They show the features of hypostatic pneumonia or broncho-pneumonia, 

without any leprosy micro-organisms being recognisable, whilst 
pneumococci and streptococci are present. 

(4) They may show signs of tubercular localisation with excavations, caseous 

pneumonia and peribronchial nodules, tubercle bacilli and without 
the leprosy micro-organisms. 

(5) In one single case he found caseous degeneration without any tubercle 

or leprosy bacilli or other bacteria. 

(6) In cases of chronic interstitial pneumonia peribronchial foci of a leprous 

nature were noticed. 

(7) The lungs may be the seat of extensive desquamative or fibrinous 

localisation of a leprous nature, with countless lepra bacilli, which 
may be accompanied by the formation of vomicae. 

(8) Gangrenous bronchial cavities, and bronchial affections of a leprous 

nature, were found in the centre of necrotic foci or surrounded by 
an interstitial or desquamative pneumonia. 

(9) Cases exist where leprous and tuberculous lesions combine, and 

accordingly are very difficult to diagnose correctly. 

The lesions of leprous lungs are very similar to those of tubercle, but it must 
be remarked that leprosy runs a more chronic course, the resulting tissues are more 
solid, caseate less frequently, and are less prone to be destroyed or eliminated. 
When considered from this standpoint leprous lesions in the lungs resemble the 
late manifestations of syphilis of the lungs. 

We are thus enabled to distinguish not only a chronic diffuse interstitial or 
trabecular form of leprosy of the lungs and a sclerotic nodular form, but also an 
acute, and at times caseating, parenchymatous variety of the disease. In the 
latter we also find fibrinous-pneumonic catarrhal lesions. 



i6 

In all cases he succeeded in proving, by means of cultures, and in some cases 
also by microscopical examination in the bronchial forms, the presence of 
diphtheroids in company with the lepra bacilli. These organisms are associated 
at times and in varying manner with bacilli of the coli group ; at other times with 
filamentary, branching micro-organisms or with streptococci or pneumococci in 
the acute pneumonic or caseating varieties (‘ Lepra ; Handbuch der pathogenen 
Micro-organismen ’ of Kolle-Wassermann, 1906, Erganzungsband, pp. 174-175).’ 

Leprotic lesions of the other organs may be accompanied by a 
greater or lesser degree of amyloid degeneration. Amyloid of the 
spleen, liver, kidneys, may be present in lepers who die with the 
features of anaesthetic cases, but typical leprotic lesions of the 
inner organs are generally only found in nodular cases. 

Liver. The most frequent change noted is a peculiar interstitial 
infiltration, accompanied by numerous acid-fast rods situated in 
‘ lepra-cells,’ that is, mononucleated round cells whose protoplasm 
is swollen out and riddled with vacuoles. The liver cells may also 
contain ‘bacilli,’ though this is not frequent. Generally the 
parenchymatous cells are but slightly altered, except for the 
presence of lardaceous disease, and under certain circumstances may 
contain a deposit of brownish-yellow pigment. The capillaries 
appear dilated and may also contain lepra-micro-organisms, which 
may have invaded the endothelium or lie loosely in the lumen. 

In common with the other organs it is often very difficult to 
decide in sections of the liver whether we are dealing with simple 
deposits of ‘ bacilli ’ which have been carried by the blood stream 
and have accumulated passively in the walls of the capillaries, or 
whether we are dealing with a chronic but progressive pathological 
process involving multiplication of the germs and an ever increasing 
local spread of the leprous lesions. 

In single cases giant cells and necrosis were detected, but I 
fear that in these instances a complicating tuberculosis could not 
be excluded. The same applies to the similar alterations seen in 
the spleen. 

Sfleen. In the spleen the acid-fast bacteria can be so numerous 
that their presence can be detected by the naked eye on staining. 
They may be scattered about in small clumps in the pulp, 
especially in the neighbourhood of the capillaries. The presence 
of lepra-cells may be under certain circumstances a noticeable 
feature of the lesion. Though a spleen may be in an advanced 




17 


stage of leprotic invasion the resulting enlargement may be hardly 
noticeable. 

Testicles . The connective tissue between the tubuli is seen to 
have proliferated actively, and to contain more or less numerous 
acid-fast rods, some lying loosely, others enclosed in lepra-cells and 
in the endothelium of capillaries. 

Numerous degrees of connective tissue proliferation can be found 
in the testicles from different patients, and under certain circum¬ 
stances even from different parts of the same testicle. The bacilli 
present also vary very much in numbers—from few bacilli, which 
can only be detected by means of an oil-immersion lens, to deposits 
rivalling in numbers those seen in the nodules of some lepers. 

Lymphatic glands . There is no doubt that the bronchial, 
mesenteric, and other lymphatic glands can be affected by a pure 
leprous infiltration, which microscopically shows very numerous 
acid-fast rods enclosed in cells of different sizes, some with 
hypertrophied protoplasm containing numerous vacuoles. Lymphatic 
glands may also be the seat of a mixed infection with tuberculosis 
and show necrosis and caseation. 

From this brief survey it will be seen that in dealing with the 
microscopical diagnosis of leprotic lesions we are continually faced 
with the difficulty of distinguishing lepra from tuberculosis, and 
that in many instances the decision must be left with the results of 
inoculation into guinea-pigs. 

It should not be forgotten that leprosy is a slower and more 
chronic complaint than even tuberculosis, and that in certain 
instances the possibility is given that a tubercle which has 
spontaneously healed may be invaded by Hansen's 4 bacillus,’ in 
which case the crucial experiment on animals would give a negative 
result, though the histological lesions might show a definite 
tuberculous character. 

With the exception of cases where the presence of numerous 
giant cells of the Langhans type, or very similar in appearance to 
those seen in tuberculosis, coupled with necrosis or caseation, 
render the presence of a mixed infection with tubercle probable, 
the main microscopical features of a leprotic lesion can be summed 
up in the following words: 

Very numerous acid-fast micro-organisms, singly or in matted 



i8 


clumps, lie loose in the tissues and lymphatics or wedged in 
compact masses in interstitial spaces. Similar acid-fast bacteria 
may be present in varying numbers in the protoplasm of various 
cells, in which under certain circumstances vacuoles can be detected. 
The endothelial cells of the lymphatics may also show a marked 
degree of bacterial invasion. The tissue reaction and production 
of connective tissue is relatively insignificant in all instances, and 
the resulting degeneration or necrosis of the tissues affected is quite 
minimal in proportion to the number of germs present in the lesions. 


REFERENCES 

Aucanazy, M. (1911). Die Leprabazillcn. Lehrbuch der path. Anatomie herausgeg. von 
Aschoff, I, 177-180. 

Jadassohn, J. (1913). Lc. 

Lit, H. P. (1911). Uber Tuberkuiose bei Leproesen. Archiv f. Dermat. u. Syphilis, CVII, 
3-16. 

MacLeod (1910). A brief survey on the present state of our knowledge of the bacteriology 
and pathological anatomy of leprosy. Lepra XI, 309-320. 

Schaeffer, J. (1900). Die Visceralerkrankung der Leprosen. Lepra I, 11-30 and II, 57-88. 


IV. MORPHOLOGY AND STAINING PROPERTIES OF 
HANSEN S ‘ BACILLUS 1 

The Mycobacterium leprae belongs to a group of acid-fast 
bacteria which in morphology and staining reactions are very 
similar to the Mycobacterium tuberculosis . The group may be 
considered to consist of the micro-organisms causing spontaneous 
tuberculous disease in human beings, cattle, horses, pigs, birds, 
etc. All these bacteria are relatively easily isolated on artificial 
media, at body heat, from the lesions in warm-blooded animals, 
with morphological and tinctorial properties analogous to those 
which may be seen in infected tissues. A sub-division of this group 
embraces several acid-fast germs, which are generally present in 
diseased tissues in enormous quantities and are extremely difficult 
to isolate in artificial culture. This sub-division may be considered 
to include the Mycobacterium leprae , the M . leprae rodentium (rat 
leprosy ‘bacillus’), and the micro-organism of Johne’s disease in 
cattle, and that causing the analogous disease of sheep mentioned 
by Twort and Ingram, Vukovic, McGowan. It is quite evident 



*9 


that a classification based on such principles cannot be satisfactory 
from a bacteriological standpoint, but it is simply brought forward 
to show once more that the leprosy ‘bacillus’ does not stand 
absolutely alone in many of its peculiarities. 

Lehmann and Neumann’s (1912) definition of the group 
Mycobacterium is the following (p. 582): 

' Thin, slender rods, often showing typical dichotomous 
branching, at times giving rise to filaments, with or without 
branching. If stained with warm carbol-fuchsine, the colouring 
matter cannot be extracted by acids, that is to say, the bacteria 
are acid-fast and behave towards stains somewhat like the spores 
of common fission-fungi. In some species the acid-fast properties 
are but slightly developed; in fact, may even be absent.’ 

According to Lehmann and Neumann’s definition, the myco¬ 
bacteria are not true * bacilli,’ and the possibility of micro-organisms 
of the group showing true branching is recognised, as also the fact 
that acid-fastness is a property which varies in different bacteria 
belonging to the group. These admissions are important in view 
of the fact that lately some bacteriologists have denied both 
conclusions when applied to the microbe of leprosy. 

A satisfactory classification of Hansen’s ‘ bacillus ’ is not a 
matter of purely theoretical interest. It has definite practical 
importance, because we have reason to expect that an exact 
knowledge of its bionomics will help isolation by means of 
artificial culture, and in such a fashion clear up the numerous 
obscure points in connection with the epidemiology and therapeutics 
of leprosy. 

The irregular morphology of the micro-organism of leprosy 
tends to confirm the view that it is not a true ‘ bacillus,’ because of 
the lack of constancy in shape, thickness and length, which in 
addition to various other features connected with the presence of 
endogenous spores, flagella, etc., determines the diagnosis 
‘ bacillus.’ The lepra bacterium is markedly pleomorphic, that is 
to say, in smears from nodules single germs can be detected which 
vary in thickness between o’3 ft and O '5 ft, and whose length may be 
anything between 1 ft and 5 ft and in rarer instances even 6 ft or 7 /a. 

Hansen’s ‘ bacillus ’ is often seen to contain small, fine granules 
(Babes-Emst) which are not as acid-proof as the rest of the 



20 


bacterium. At times, peculiar hyaline breaks can be detected in 
the rods; they have been considered to be spores. 

Another peculiarity of the bacillus is its breaking up into small 
coccoid bodies, which Deycke considered to be a sign of degenera¬ 
tion, but which the observations of Martinez-Santamaria seem to 
indicate are possibly developmental stages prior to multiplication. 

The reasons for which it is possible to classify Hansen’s 
'bacillus’ as the acid-fast stage of a Mycobacterium closely allied 
to the Actinomyces are based on morphological grounds and 
deductions by analogy, in addition to the interpretation of 
cultural attempts. 

The morphology of Hansen’s ' bacillus,’ when examined with 
high-power lenses (say 2 mm. apochromatic, comp, eyepiece No. 12) 
and in sufficiently thin smears, can be seen to be exceptionally 
variable; structures with one end tapering and the other definitely 
thickened (clubbed) are mixed with rods of uniform thickness but of 
different lengths and fragments, which seem to consist of a series 
or chain of small beadlike coccoid bacteria. 

At times in smears taken from nodules branching forms arc 
seen; they are, however, rare. These peculiar pleomorphic 
appearances, especially the clubbed and branching forms, are 
typical of the fragments of actinomycotic germs having the 
tendency to break up into acid-fast fragments. The variations in 
morphology have lately been commented upon by Galli-Valerio, 
who, in discussing the relationship of the genera Corynebacterium, 
Mycobacterium and Actinomyces, has not failed to note the 
numerous points of contact between these groups. 

From the standpoint of analogy we know from the observations 
of Petrone (1884), Metschnikoff (1888), Fischel (1892), Babes and 
Levaditi (1897), Schulze (1899), Lubarsch (1899), Lehmann and 
Neumann (1912), Galli-Valerio (1912), Foulerton, and others, that 
true branching can occur in Mycobacterium tuberculosis and that its 
appearances in sputum and tissues of experimental animals are at 
times distinctly like those brought about by actinomycotic germs, 
especially in the production of radiary deposits of bacteria. Abbot 
and Gildersleeve have noted similar appearances in connection with 
‘ saprophytic ’ micro-organisms. Further, observations of Silber- 
schmidt (1899), Galli-Valerio (1910), Birt and Leishman, and 



21 


others, have made us acquainted with a group of germs which clearly 
can be classed with the Actinomyces or Streptothrix class, yet in 
artificial cultures and on injection into animals show numerous acid- 
fast fragments very similar to M. tuberculosis. 

That an acid-fast rod seen in tissues, can be found to be a 
filamentary branching micro-organism in artificial cultures, is an 
observation which has been made so repeatedly by those working 
with A. caprae , Birt and Leishman’s Streptothrix, that there is no 
necessity to dilate further on the subject. 

Accordingly we may deduce that it is by no means far-fetched 
to suggest that Hansen’s ‘bacillus’ may present a filamentary, 
acid-labile appearance in artificial culture, though the M. tuber¬ 
culosis is consistently acid-fast on the usual laboratory media: and 
even in this instance Frei and Pokschischewsky, and later Wherry, 
have shown that under certain circumstances this property can be 
made to vary. 

The interpretation of the cultural results of investigators whose 
bacteriological technique was not open to criticism (Beauchamp 
Williams) shows us that in the great majority of cases their culture 
tubes have either remained sterile for months and months (Fraser 
and Fletcher), or that their repeated efforts were only rewarded by 
the isolation of a singularly pleomorphic diphtheroid with slight 
acid-resisting properties, that is to say, that after prolonged 
staining with carbol-fuchsine it could be, and was, only partially 
bleached by a weak acid such as one per cent, sulphuric. This 
diphtheroid was usually found to have peculiar filamentary forms. 

The relationship of the genus Corynebacteria to the Actinomyces 
is also very close; to give an example out of a numerous series we 
have the Corynebacterium ttecrophorum (Lehman and Neumann) 
which Schmorl considers as a Streptothrix and other authors as an 
Actinomyces. 

As a matter of fact, the bacteriological literature of the last ten 
years contains numerous, indeed very numerous, hints that a strict 
separation of bacteria on cultural characteristics alone is extremely 
difficult, to say the very least, because of the existence of very many 
micro-organisms which do not conform in every case with any 
single distinctive test or feature. 

It will be seen that, though definite reasons are extant to 



22 


postulate that Hansen’s ‘ bacillus * belongs in reality to the Actino¬ 
myces, Actinobacteria or ray-fungus group, still the last conclusive 
test is a matter of surmounting the difficulties inherent to the 
artificial isolation and cultivation of this bacterium. 

Staining Properties of Hansen’s ‘ Bacillus.’ 

In a recently published study on the bacteriology of human and 
rat leprosy Wolbach and Honeij (1914) come to the conclusion that: 

‘ The possibility that the non-acid-fast and acid-fast organisms isolated from 
cases of leprosy are related and may be converted one into the other is one we 
must take, however reluctantly, into consideration, because of the great length of 
time the leprosy bacillus resides in the human body and the variety of conditions 
it is subjected to by virtue of the stage of the lesion and anatomical situation. 
The loss and acquisition of properties as distinctive as that of acid-resistance are 
well known in the field of bacteriology and indeed the changes undergone by the 
tubercle bacillus towards staining reactions is the strongest claim for the possibility. 

Very numerous methods have been published with the purpose 
of communicating a strictly specific stain for the leprosy micro¬ 
organism, that is, a stain which would show up in some distinctive 
colour Hansen’s 4 bacillus,’ whilst all other bacteria, however similar 
but not identical, would not take the same hue. The majority of 
these methods are based on the property of the M. leprae of taking 
up carbol-fuchsine or a similar stain more readily than Koch’s 
1 bacillus,’ and, moreover, on the greater resistance, as a rule, of the 
latter to the bleaching properties of mineral acids or strong alkalis. 

Accordingly it will be seen that we are dealing with differences 
in degree and not in quality, and as to be expected, such methods 
of differentiation are apt to fail in a fair proportion of cases and be 
unreliable in the remaining instances. 

Our experience in dealing with the sections and smears in the 
laboratory at Robben Island is that to obtain well stained leprosy 
bacilli in sections it is advisable to use 2 \ per cent, carbolic in 
making up the fuchsine solution, to stain in the warm solution one 
or two hours according to thickness of section, and to differentiate 
with one per cent, hydrochloric acid in the common methylated 
spirit of commerce. Counterstain w'ith alum-haematoxylin. 

Skin fragments are always fixed in absolute alcohol, specimens 
from organs in 4 per cent, formaldehyde. Embedding in paraffin 
is advisable, though excellent sections can also be got in celloidin. 



23 


For the quick staining of nasal smears, etc., Pappenheim’s 
carbol-fuchsine-methylene-blue-coralline method gives reliable 
results. 

Unna has described a method of distinguishing lepra ‘bacilli’ 
which have been fixed alive from those that were brought dead into 
the fluid. The validity of this distinction is, however, only founded 
on circumstantial evidence. 


REFERENCES 

Foulerton, A. G. R. (1910). The Milroy Lectures on the Streptothrichoses and Tuberculosis. 
Lancet, I, 551. 

Frei und Pokschischewsky (1911). Centralbl. f. Bakt., LX, p. i6r. 

Lehmann und Neumann (1912). Atlas und Grundriss der Bakteriologie. 

Wherry (1913). Some chemical conditions influencing acid-proofness and non-acid-proofness 
in a saprophytic culture of B. tuberculosis. Joum. of Inf. Dis., XIII, 144-154. 


V. ARTIFICIAL CULTIVATION OF THE MICRO-ORGANISM 

OF LEPROSY 

The first attempts to cultivate Hansen’s “ bacillus ” were made by Sir Patrick 
Manson many years ago. at a time when aniline dyes had not yet been introduced 
into bacteriological technique. Later on. in 1882, Hansen thought he had 
succeeded in getting the “ bacillus ” to grow artificially in some serum contained 
between a microscope slide and slip, but he never followed the matter any further 
after mentioning this observation. 

Neisser in 1886 considered that multiplication of “bacilli” from a nodule 
had taken place on cooked egg and blood serum. In any case he was not able to 
get further generations to grow. 

Bordoni-Uffreduzzi in 1887 cultivated from the bone marrow of a case of 
leprosy a partially acid-fast or acid-resistant micro-organism, which, from the 
description given and the specimens seen. I would consider to be morphologically 
a diphtheroid. It was not quite as acid-fast as the Bacillus leprae in tissues. 

Campana from 1889 onwards has described micro-organisms, which he 
cultivated anaerobically from the nodules of lepers. I have been able to examine 
the culture : it is a mixture of cocci, and coli-like organisms, none of which are 
add-fast or even acid-resisting. There appears to be a sad lack of technique in 
isolating these cultures, an opinion also apparently shared by Wasserman. 

Kanthack and Barclay, in 1891, cultivated a micro-organism of varying add- 
fast properties from a leproma. Originally they considered it to be leprosy,, but 
later on they withdrew their claim. 

Levy in 1898 cultivated from a leper a slightly add-fast, filamentary organism, 
which in later publications he considered identical to that isolated by Czaplewski 
in the same year from another case of leprosy. No animal experiments succeeded 
with these micro-organisms. 

Spronk. also in 1898, was successful in isolating a diphtheroid from lepromas. 
He carried out some agglutination experiments with it, which, however, appear 



2 4 


not to have had conclusive results owing to incomplete knowledge of the 
agglutinating properties of sera at that time. 

Babes in 1899 isolated on several occasions, from nodules and the inner organs 
of lepers, an acid-resisting diphtheroid, illustrations of which he has given, and 
which I consider extremely suggestive. Unfortunately he did not follow the 
work up, as he later considered that these diphtheroids were contaminations. 

Barannikow in 1899 cultivated also a diphtheroid from lepers, very similar to 
the one isolated by Bordoni-Uffreduzzi, Spronk, Babes, Levy, Czaplewsky. This 
he considered was a pleomorphic stage of Hansen’s “ bacillus.” 

Kedrowsky published his first results in 1900. Using placental-extract-agar, 
he succeeded in isolating from the nodules of lepers an extremely pleomorphic 
micro-organism, with varying acid-resisting properties. 

Weil in 1906 succeeded in getting what he thought was a multiplication of 
“ bacilli ” by injecting leper serum into a hen’s egg, which he incubated at 37°C. 
He was not able to carry this further than one, the first, generation. 

Rost published his first results in 1904. He made a great point of the fact 
that the least trace of salt would not allow the culture to succeed. The medium 
used was the extract from rotten fish. Semple in 1905 was sent to confirm Rost’s 
results and found that his isolations were due to faulty bacteriological technique. 
This was also the opinion of other bacteriologists who had occasion to examine 
his work. 

Rost’s culture is a dark red, wrinkled, dry culture. It is markedly chromogenic, 
especially on gelatine at room temperature, under which conditions it grows quite 
easily. At 37 0 C. it grows rapidly on most of the usual culture media, sometimes 
within 48 hours. 

Morphologically it is a slightly acid-resisting streptothrix, with add-fast 
fragments ; this property varies, however, in different culture tubes. It is Gram¬ 
positive. 

It resembles Grassbergeris isolation from butter in culture and under the 
microscope. It has not produced leprotic lesions in animals. 

Shiga (1909) isolated on potato-serum-glycerine a pleomorphic, partially 
acid-resisting diphtheroid. 

Clegg in 1909 published his results and concludes with the following summary : 

1. The leprosy bacillus was first cultivated from leprous material in 
symbiosis with other unidentified bacteria and amoeba, and later from 
other cases in symbiosis with amoeba and cholera vibrion. 

2. By heating a symbiotic culture of amoeba, cholera, and leprosy for 
half an hour at 6o°C. and incubating, the leprosy barillus was obtained in 
pure culture. 

3. The leprosy bacillus, isolated in this manner, grows readily on the 
ordinary laboratory culture media. 

4. The bacillus is pathogenic for guinea-pigs, subcutaneous inoculations 
having caused lesions, which macroscopically and microscopically resemble 
the leprous lesions of the human subjects. 

Clegg’s culture is granular, dry, yellow, grows at room temperature on gelatine 
or any of the common laboratory media. I have found difficulty in getting the 
strain to grow in the incubator. It is add-fast, alcohol-fast, and Gram-positive. 
Morphologically it consists of short plump rods, very similar in appearance to the 
timothy grass bacillus ; at times, however, it may resemble the ‘ 4 Smegma bacillus,” 
and accordingly consist of slender matted rods. 



25 


On injection into animals it does not produce lesions comparable with leprosy 
in man. 

Currie, Brinckerhoff and Hollmann in 1910 succeeded after several failures, 
in growing acid-fast rods in symbiosis with cholera vibriones and amoebae. They 
divided up their cultures into four “ strains,” and found that one was practically 
identical with Clegg’s culture from Manila. 

Serra (1910) cultivated anaerobically from several cases (three of seven 
attempted) a peculiar micro-organism which is Gram-positive, but not acid-fast, 
as it can be bleached by 2 per cent, nitric acid. Morphologically it appears to 
be a “ diphtheroid.” 

Twort in 1910 by using his “ ericoline ” method isolated from the nasal 
discharge of a leper, on egg-medium to which he had added ground-up tubercle 
bacilli, an acid-fast rod, which grew very slowly indeed, and was morphologically 
identical with the “ bacilli ” found in tissues. He was not able to carry the culture 
further than the first generation, but still his incomplete results deserve considera¬ 
tion, in view of his success with Johne’s “ bacillus.” 

Duval in 1910 and the following years published various results in connection 
with the isolation of Hansen’s “ bacillus.” 

His first publication was founded on Clegg’s results, and the isolation of the 
acid-fast micro-organisms was made with amoebae in the fashion identical with that 
of Clegg. The micro-organism in question appeared, however, to differ from 
that isolated by the latter. In later publications Duval found he could dispense 
with the amoebae and cultivate the “ bacilli ” from the nasal discharge of lepers 
simply by adding amino-acids to the culture media. Later on, even the addition 
of the amino-acids does not seem to be indispensable, for an isolation is mentioned 
on Novy-Mac Neal’s blood-agar. The last papers deal with isolation by means 
of Kedrowsky’s placental juice agar, which had been re-discovered by Wellman 
and published as “ Wellman’s placental agar.” Animal experiments failed to 
produce typical leprous lesions. 

Duval’s so-called culture of lepra is a bright yellow, moist, smooth culture. 
It grows easily and abundantly on gelatine by room temperature and can be easily 
regained in pure culture on common agar from the organs of animals that have 
been inoculated with it. At 37 0 C. it takes about three days to grow on any of 
the current laboratory media. 

Morphologically it is coccoid in appearance, but under circumstances, when 
injected into animals and sometimes on artificial media, it is apt to present a 
somewhat more elongated appearance, but at no time closely resembles Hansen’s 
“ bacillus.” 

It is acid-fast and alcohol-fast. Gram-positive. 

Its cultural and morphological characteristics cause it to resemble markedly 
a * saprophytic acid-fast micro-organism isolated by Nabarro from London milk. 
Therefore it is not astonishing that animal experiments did not succeed in 
producing any lesions analogous to those found in lepers, and that at the 
International Medical Congress in London (1913) this author admitted he had 
no culture of leprosy to show or hand round. 

Kedrowsky’s paper published in 1910 deserves special mention in view of 
the care with which the experiments have been carried out. They took ten years 
to complete. 

His first isolations from the nodules of three lepers gave as a result two distinct 
bacteria; one was a non-acid-fast filamentary, interlacing, branching micro¬ 
organism, the other was a slightly acid-resisting diphtheroid. He also cultivated 



26 


other micro-organisms, which, however, from his pictures and descriptions are 
easily seen to be but variations of these two groups. He injected these various 
bacteria strains singly, into mice and rabbits, in different fashions (under the dura, 
intravenously and intraperitoneally) and observed the animals for prolonged 
periods, in some cases over two years. He found that, whatever micro-organism 
he had injected, the result was practically the same in all cases; very numerous 
acid-fast micro-organisms of the “ tubercle bacillus ” type in the viscera, from 
which they could be regained in pure culture as acid-fast rods. The resulting 
lesions resembled in some cases the chronic type of tuberculosis induced in rabbits 
by injections of human tuberculosis ; in others the lesions were strikingly similar 
to those occurring in visceral leprosy of human beings. 

Kedrowsky’s acid-fast culture of “Hansen’s Bacillus” is a moist, creamy, 
smooth or wrinkled ivory-white culture which resembles avian tuberculosis to a 
very marked extent. It grows only at incubator temperature on special media, 
such as placental-juice-agar, glycerine-agar, horse-serum-nutrose-agar or any 
similar medium suitable for tubercle. Multiplication is generally apparent within 
ten to fourteen days, but may take three weeks to four weeks or more to attain 
its maximum. 

No growth takes place at room temperature or on gelatine. 

It is add-fast, cannot be bleached by 20 per cent, nitric add in one minute 
after staining for five minutes with warm carbol-fuchsine. It is alcohol-fast, will 
withstand absolute alcohol for ten minutes. Gram-positive. Morphologically it 
is markedly pleomorphic, and resembles avian tuberculosis in this respect, in fact 
I do not believe that these two bacteria can be distinguished from one another 
under the microscope, except that Kedrowsky’s strain of lepra is more apt to show 
clubbed shapes than even avian tubercle. 

Williams in 1911 isolated diphtheroids and streptothrices from the nodules 
of lepers. He considered them to be pleomorphic stages of Hansen’s “ bacillus,” 
but was not able to produce any typical lesions in animals with the micro-organisms 
isolated. 

Currie, Clegg and Hollmann express their opinion on these cultures in the 
following terms:— 

“ This investigator considers that his work tends to confirm the work of Rost, 
Deycke, and Clegg, i.e., that all these investigators have, by different methods, 
isolated the same organism, which latter is very pleomorphic. He seems, however, 
to base this opinion on very little data other than that the organisms in question 
were all grown from lepromata, and that local nodules were produced in animals 
by inoculation of some of these cultures.” 

Bayon (1911) published the first confirmatory results of Kedrowsky’s isolations, 
above all in relation to the existence of filamentary non-acid-fast micro-organisms 
in cultures taken with all aseptic precautions from leprous nodules. He also 
confirmed the possibility of isolating on various media diphtheroids which were 
slightly acid resisting. Both bacteria acquired acid-fast properties after injection 
into animals. This author considered that the identification of bacteria isolated 
from lepers was the most important point in connection with the bacteriological 
work, and accordingly carried out extensive investigations by means of numerous 
experiments on animals, serological tests and observations on human beings. He 
found that Kedrowsky’s view that the parent form of Hansen’s “ bacillus ” is a 
filamentary, interlacing, branching, non-acid-fast micro-organism is correct, and 
that this pleomorphic property is shared by several other acid-fast bacteria, e.g. ; 
the smegma “ bacillus,” some saprophytes, such as the acid-fast rod found in 
tap-water, etc. His comparative experiments with Duval’s. Rost’s, Clegg’s cultures 



27 


persuaded him that they behave in culture, on common gelatine at room tempera¬ 
ture, and in animal experiments, like the usual ubiquitous, acid-fast micro-organisms 
found in grass, dung, milk, butter, earth, etc. This observation he confirmed by 
serological tests, which showed that these cultures when used as antigens do not 
react specifically with any lepers’ serum. The lesions produced in animals are 
not like the lesions of rat or human leprosy. On the other hand, he found that, 
given sufficient time for incubation, and notwithstanding a very heavy percentage 
of failures, Kedrowsky’s strain was capable of producing in rats, mice and rabbits 
lesions extremely similar to those occurring in visceral lepra of human beings. To 
obviate as much as possible any experimental error, the culture was injected into 
guinea-pigs and fowls to determine whether, as Babes had suggested, it might not 
be a tubercle strain which had infected Kedrowsky’s animals. Small doses (five 
to ten loopfuls) did not cause any lesions in guinea-pigs or fowls, even after pro¬ 
longed observation. 

The serological tests employed also showed this strain to be capable of reacting 
specifically with the serum of certain lepers. A filtered cultural extract when 
injected into lepers caused a rise of temperature to take place. This reaction, 
however, has no specific value, as it can be brought about by many different sub¬ 
stances, and is, moreover, not constant. 

Bayon’s culture (in the diphtheroid stage) was studied by Priestley, who com¬ 
pared it with numerous (namely, 48) other micro-organisms from various sources, 
showing somewhat similar morphological features, with the intention of bringing 
evidence to bear on the point, whether this diphtheroid could possibly be an 
ubiquitous saprophyte, whose presence in the skin of lepers had only casuistic, 
and not etiological importance. 

Priestley found that morphologically the “ bacilli ” were very irregular in 
shape, clubbing was very marked, polar granules could be detected by Neisser’s 
stain. Gram-positive. On serum they formed raised, white circular colonies, 
like Klebs-Loeffier “ bacillus ” ; on agar, very minute, circular, greyish-white 
colonies; no growth on gelatine; on broth, very slight growth, very stringy 
deposit; no growth on potato. Non-virulent. They fermented dextrose and 
laevulose, but not saccharose, lactose, maltose, mannite, dulcite, glycerine, dextrite, 
galactose, did not clot milk. Unfortunately the study was not completed at the 
time, and no further details were published because all attention had to be con¬ 
centrated in attempting to bring the peculiar results of some American authors 
in relation with the observations made on Kedrowsky’s culture and the acid-fast 
micro-organism isolated from rats inoculated with acid-labile cultures from lepers. 

Numerous experiments on animals showed, however, that the acid-fast stage 
of the filamentary micro-organism was capable of producing in rabbits, rats and 
mice lesions very similar to those obtained by Kedrowsky in his experiments, and 
which accordingly were analogous to the lesions observed in leprosy of the human 
being. 

The result of these numerous experiments, prolonged over several years (4), 
was accordingly that Kedrowsky’s isolation was true leprosy, and that the cultures 
of some of the other authors appear to be accidental. 

Reenstierna (1913) succeeded in cultivating from the blood and nodule of a 
leper an acid-fast rod, which was morphologically similar to Hansen’s “ bacillus ” 
seen in tissues, and numerous acid-labile bacteria. The acid-fast rods were 
isolated in pure culture by the means of 10 per cent, antiformin, and lasted four 
generations in their add-fast condition, and then turned acid-labile. 

A pure culture isolated from these acid-labile bacteria showed on single 
occasions the presence of acid-fast micro-organisms. 



28 


A monkey (Macacus rhesus) was injected in the brain and ischiatic nerve with 
a pure culture of the acid-fast rod isolated from the blood of a leper. This animal 
showed peculiar maculae on the chest and face, which contained acid-fast micro¬ 
organisms. The death of the monkey took place three months after injection, 
and the inner organs showed numerous caseating nodules which contained big 
bundles of acid-fast rods, which were mostly situated extra-cellularly. 

Another Macacus rhesus was injected with a pure culture of the acid-labile 
micro-organism isolated from the blood of a leper. Forty-two days after, it 
developed blisters on its fingers and toes, which contained acid-labile and acid-fast 
bacteria. Shortly after the animal was killed, and acid-fast rods were found at 
the site of injection, and in caseating inguinal and sacral lymphatic glands. 

A third M. rhesus was injected with a culture of the acid-labile micro-organism, 
which was isolated from a single bacterium by means of Bum’s Indian ink method. 
This animal lived eight months, and though during its life it developed a spot 
which contained diphtheroids and a contracture, the autopsy revealed no leprotic 
lesions. 

Reenstierna concludes from his numerous observations that the bacillus of 
leprosy is not only morphologically but also biologically closely related to the 
tubercle “ bacillus.” 

His experiments deserve special consideration, because they belong to the few 
which have been undertaken without the preconceived idSe fixe that leprosy and 
tubercle are absolutely distinct diseases in every single particular, and that Hansen’s 
“ bacillus ” is acid-fast from beginning to end of its existence, notwithstanding 
that there is sufficient evidence to show that either statement need go unchallenged. 

Stanziale (1913) cultivated from the eye of a rabbit, which had been inoculated 
128 days before with a fragment of leprotic nodule under anaerobic conditions, on 
agar-egg-yolk, an acid-fast rod very similar to that of tuberculosis. The original 
culture took a month to develop, but on other media the colonies were already 
apparent after twenty-four to forty-eight hours. On agar and broth the culture 
resembles tubercle, except for the fact that it is more abundant. Apparently the 
resistance to the bleaching properties of acids varies, though the bacterium contains 
granules which are markedly acid-fast. Gram-positive. Numerous clubbed 
shapes similar to those seen in Corynehacterium diphtheriae. Non-pathogenic for 
animals. 

Wolbach and Honeij (1914) have lately isolated a diphtheroid from a case of 
leprosy. It was originally cultivated aerobically on ascitic fluid dextrose agar, 
and appeared within ten days in the form of a translucent whitish band around 
the piece of gland tissue. 

“ When first isolated the growth was poor on bouillon, with or without glycerin, 
and on plain agar, dextrose agar and glycerin agar. After several months’ cultiva¬ 
tion the bacillus grew readily on these media. No growth has been obtained 
upon potato, with or without the addition of 0*5 to 1 per cent, sodium carbonate. 
Glycerin does not favour the growth of this bacillus.” Gram-positive. After 
staining with carbol-fuchsine (how long ?) they resisted decolourization with 1 per 
cent, hydrochloric acid in 70 per cent, alcohol; 20 per cent, sulphuric acid left the 
bacilli from litmus milk distinctly red, after staining by Gabbett’s method for 
tubercle. No lesions were obtained on inoculation into Japanese waltzing mice, 
guinea-pigs, rabbits and white rats. The methods of inoculation were intra¬ 
venous, subcutaneous and intraperitoneal. 

Wolbach and Honeij consider this culture to be identical with the diphtheroid 
which has so often been isolated from cases of leprosy, and put in a plea for more 
full data about any micro-organisms which may be cultivated in future. 



2 9 


These are the results obtained in the course of upwards of thirty 
years of bacteriological investigation of an extremely widely spread 
disease. No doubt hundreds of negative results have been obtained 
and not published, but it can be said that as a rule every time the 
problem has been tackled by a competent bacteriologist the culture 
and isolation of a diphtheroid or filamentary germ has been the 
result. This has also apparently been the case with the investiga¬ 
tions of Fraser and Fletcher, who, however, discarded all 
diphtheroids because of their ubiquity, a standpoint which would 
considerably simplify pathology. The micro-organisms of syphilis, 
of tuberculosis, of cholera, could also, on similar grounds, simply 
be ruled out of court, as far as their etiological significance is 
concerned. 

The main point has, however, always been that as a rule 
these diphtheroids, though slightly or partially acid-resisting, 
did not turn absolutely acid-fast on injection into animals, with the 
exception of the instances mentioned. 

It should be understood that every experiment connected with 
leprosy should be repeated scores of times, because evidently if the 
matter were simple it would have been solved from the very 
beginning. The crucial experiment to be carried out with any 
culture isolated from a leper is not only to get it to acquire the 
acid-fast properties and the morphology of the ‘ bacillus * seen in 
tissues, but also to succeed in producing lesions analogous to those 
seen in lepetS and in rats spontaneously infected with M. leprae 
tnurts. Bearing in mind the great difficulties encountered in 
transmitting leprosy to animals through the injection or inoculation 
of nodules teeming with ‘ bacilli/ we must be prepared to face series 
and series of negative results. As negative results, however 
numerous, are powerless to destroy a single positive observation, it 
must be admitted that Kedrowsky has published micro-photographs 
and drawings of lesions in rabbits and mice which are strikingly 
like those produced by the inoculation of human and rat * virus 1 in 
rabbits and rats. Such specimens have not been published by any 
other author, excepting those to be found at the end of this paper; 
the corresponding deduction should not be difficult. (Plate II, 
fig. no 



3 ° 


REFERENCES 


(Arranged in chronological order.) 


Nusser, A. (1886). Histologische und bakterioiogische Lepra-unter*uchungen. Virchow’* 
Archiv, CIII, 355. 

Bordoni-Uffriduzzi, G. (1887). Uber die Kultur der Leprabazillen. Zeit*chrift fur Hygiene, 
III, 178. 

Manson, Sir P. (1890). Journal of the Leprosy Investigation Committee, I, 40. 

Stallard, J. H. (1890). The Lepra Bacillus. Occidental Medical Times, IV, 220. 

Campana, R. (1891). Un badllo simile al badllo leproso sviluppatosi in tentativi di cultura di 
tessuti con lepra tuberculare. Riforma medica, No. 14. 

Ducrey, A. (1892). Tentativi di cultura del badllo della lepra con risultato positive. Giornale 
italiano delle malattie veneree, etc. 

Levy, E. (1898). Ein neues, aus cinem FaBe von Lepra gezfichtetes Bakterium a us der Klasse 
der Tuberkelbadllen. Archiv ffir Hygiene, XXX, 168. 

CzAPLEWsrr, E. (1898). Uber einen neuen, aus einem Leprafalle gezfichteten, alkohol- und 
sSurefesten Bacillus aus der Tuberkelbadllengruppe. Centralblatt ffir Bacterio logic, 
XXm, 97 and 98. 

Spronk, C. H. H. (1898). De cultur van de badl van Hansen en de serodiagnostik van lepra. 
Nederl. Tijdschr. van Geneeskunde, II, 322. 

Teich, M. (1899). Beitrage zur Kultur des Leprabadllus. Centralblatt fur Bakteriologie, 
XXV, 756. 

Babes, V. (18^9). Uber die Kultur der von mir gefundenen Diphtherideen. Centralblatt fur 
Baktenologie, XXV, 12$. 

Barannikow, J. (1899). Zur Frage uber die Bakteriologie der Lcpromata (Vorlaufige 
Mittheilung). Centralblatt ffir Bakteriologie, XXV, 113. 

Kedrowsky, W. J. (1901). Beobachtungen fiber die kfinstliche Kultur des Lepraerrcgers. 
Zeitschrift ffir Hygiene, XXXVII, $2. 

-(1904)* Experiraentelle Erfahrungen fiber Lepraimpfungen bei Tieren. Centralbl. 

ffir Bakteriologie, XXV, 368. 

- (1910). Experimentelle Untersuchungen fiber Lepraimpf ungen bei Tieren. 

Zeitschrift fur Hygiene, LVT, 1. 

Van Houtum, G. (1902). A successful attempt to cultivate Bacillus leprae. Journal of 
Pathology, VIII, 260. 

Zenoni, C. (1902). Ricerche batteriologiche sulla lebbra. Gazzetta medica italiana, LIII, 
No. 45 - 

Levy, F. (1902). La coltura artifidale del bacillo della lepra. Giornale della Sodeta italiana 
d’lgicne, XXIV, 219. 

Gjubert, J. (1903). Zur Bakteriologie der Lepra. Baumgarten’s Jahresbericht, XIX, 337. 

Karlinsky (1903). Kulturen eines aus Lcprabaciilen gezuchteten saurefesten Bacillus. 
Monatshefte ffir praktische Dermatologie, XXXVII, No. 9. 

Rost, E. R. (1905). The cultivation of the Bacillus leprae. Indian Medical Gazette, February. 

Deycke und Reschad (1905). Neue Gesichtspunkte in der Leprafrage. Deutsche mcd. 
Wochenschrift, No. 13, p. 489. No. 14, p. $45. 

Weil, E. (1906). Essais de culture du bacillus lepreux. Annales de l’lnstitut Pasteur, 1905, 
793 - 

Clegg, M. T. (1909). Some experiments on the cultivation of Bacillus leprae. Philippine 
Journal of Sdence, IV, 77. 

-(1909). The cultivation of the leprosy bacillus. Ibidem, IV, 403. 

Currie, Brinckerhoff* and Hollman (1910). On the cultivation of the bacillus of leprosy 
by the method of Clegg. Public Health Reports, page 1173. 



3i 


Duval, C. W. (1910). The cultivation of the leprosy bacillus and the experimental production 
of leprosy in the Japanese dancing mouse. Journal of Experimental Medicine, XII, 649. 

- (1911). The cultivation of the leprosy bacillus from the human tissues, with special 

reference to the amino acids as culture media. Journal of Experimental Medicine, 
XIII, 365. 

Twort, F. (1910). A method of isolating and growing the lepra bacilli of man. Proceedings 
of the Royal Society, LXXXIII, 156. 

Skr&a, A. (1910). Contributo alio studio del bacillo di Hansen. Giora. ital. Mai. ven. Fasc. II. 

-(1911)* Inoculation de culture du bacilli de Hansen dans l’oeil du lapin. Lepra, 

XII, 1-14. 

Williams, T. S. B. (1911). Leprosy: A new view of its bacteriology and treatment. 
Supplement to the Indian Medical Gazette, May, p. 1. 

Bayon, H. (1911). Demonstrations of specimens relating to the culture of the leprosy bacillus. 
British Medical Journal, November nth. 

- (1912). A comparative experimental study of the leprosy cultures of Clegg, Duval, 

Kedrowsky, Rost and Williams. British Medical Journal, November 2nd. 

- (1914). The Micro-organism of leprosy, has it been cultivated ? Lepra, XIV, 187-192. 

Liston, W. G. and Williams, T. S. B. (1912). A Streptothrix isolated from the spleen of a 
leper. No. 51, Scientific memoirs (Med. and Sanitary Depts.) Govt, of India. 

Rzenstieina, J. (1913). Uber die Kultivierbarkeit und Morphologic des Lepra-erregers und 
die Ubertragung der Lepra auf affen. Archiv fur Dermatologie, CXVI, 1-76. 

Stanzialx, R. (1913). Isolamento colturale di un bacillo acido resistente da occhio leproso di 
coniglio. Atti della R. Accademia Napoli, No. 2, pp. 3-7. 

Wolbach and Honuj (1914). The diphtheroid bacillus from leprosy lesions. Journal of Med. 
Research, XXX, pp. 1-8. 


VI. THE EXPERIMENTAL TRANSMISSION OF LEPROSY 

TO ANIMALS 

Leprosy is definitely known to be a bacterial disease, and 
therefore it should not and cannot be impossible to transmit it to 
animals by experimental inoculation, if it is considered that practi¬ 
cally all pathogenic bacteria can be made to produce in animals 
diseases similar or analogous to those caused in man. In fact, the 
existence of a disease in rats, extremely similar to leprosy, ought 
to throw light on this vexed question and postulate the possibility 
of the experimental infection of laboratory animals. 

In estimating the value of any experiments made for the purpose 
of inoculating animals with leprosy, it should be remembered that 
but fourteen years ago it was universally denied that syphilis could 
be given to rabbits or monkeys. Even scientists like A. Neisser 
upheld this belief, which, as we now know’, was absolutely 
erroneous and totally unfounded. Syphilis is extremely contagious, 
develops comparatively rapidly, therefore it is not astonishing that 



32 


the transmission to animals should be a relatively easy matter, much 
more so than leprosy. Yet, even after the communication of the 
first successful inoculations of syphilis to rabbits and monkeys, 
numerous negative experiments were published, accompanied by 
the statement that the sores produced on the testis of rabbits, etc., 
were not produced by the syphilitic virus but were due to the effects 
of contaminating bacteria. 

These recent happenings should induce more attention being paid 
to the evidence which has accumulated in the course of time, 
showing that though in the great majority of cases leprotic 
products, when inoculated into animals, are eliminated without 
leaving any definite traces, still multiplication and conclusive 
dissemination of the bacilli does take place in a small number of 
instances. 

In 1881, Neisser, after twenty-four unsuccessful attempts on rabbits, 
injected two dogs with leprous nodules, and concluded from the results that leprosy 
developed locally at the site of inoculation. 

1883. Damsch grafted lepromata in the anterior eye chamber of two rabbits. 
The animals died after 139 and 219 days. Apparently a slight increase in the 
quantity of acid-fast germs had taken place. He also inoculated two cats in the 
abdomen with leprous nodules. After 120 days numerous bacilli were found in 
the newly-formed tissue round the atrophied nodules. 

1888. Vossius carried out experiments in a similar fashion to Damsch on 
rabbits and had identical results. He also considered that the acid-fast germs had 
multiplied. 

1885-6. Melcher and Ortmann inoculated four rabbits in the same way. 
They succeeded in getting, after four to ten months, definite metastases and deposits 
of acid-fast germs in the spleen, liver, caecum, pleura, and pericardium. They 
stained these germs with Baumgarten’s differential stain, and found that they had 
the staining properties of Hansen’s “ bacillus.” 

1887. Wesener injected several rabbits with leprous nodules, and in two 
animals out of eight, after periods of four and a half months and eight months 
respectively, he found lesions apparently identical with those described by Melcher 
and Ortmann—that is, nodules from the size of a pinhead to that of a pea in the 
lungs, on the pleura, on the epiploon, in the liver, in the lymphatic glands, spleen, 
kidneys, caecum, and peritoneal surface. Caseation and giant cells were present. 
In the eyes of the rabbits he obtained after six and eight months a congregation of 
round cells choked up with acid-fast micro-organisms. He considered that the 
generalised lesions were due to tuberculosis, and that the ocular lesions were caused 
by dead bacilli, because acid-fast germs can be demonstrated in the eye of the 
rabbit after inoculation with nodules which have been kept for years in alcohol. 

1901. Barannikow confirmed these results on one rabbit. 

1893. Wnoukow inoculated twenty rabbits with leprous nodules in various 
fashions (intraocularly, subcutaneously, intraperitoneally). In fourteen rabbits 
he got lesions which he considered to be of tuberculous nature. No confirmatory 
tests with guinea-pigs appear to have been made. 



33 


1893. Tedeschi inoculated a leproma into the dura of a monkey. The 
animal died after eight days; that is far too brief a period for any conclusions to 
be drawn. 

1902. Ivanow injected several guinea-pigs with leprous nodules. In one 
case the animal was killed after eight months and found to have nodules in the 
epiploon, which Ivanow considered were due to multiplication of the bacilli. 

1905. Thiroux inoculated five rabbits with leprous material. The animals 
lived thirteen to twenty months, and presented various lesions at autopsy, which 
Thiroux considered to be tuberculous, after positive inoculation of two guinea- 
pigs. 

1906. Nicolle inoculated monkeys with ground-up nodules, and succeeded 
in getting localised lesions. 

1909. Marchoux and Bourret inoculated a chimpanzee under the skin of 
the ear with a freshly-excised leprous nodule. It apparently increased in size 
during three months, and then began to be resorbed. An examination of the 
blood showed a few leucocytes with badly-staining bacilli. The animal died ninety- 
six days after inoculation. The nodule had reached the size of a split pea. and under 
the microscope was shown to consist of three layers. In the middle were the 
remains of the original tissue, quite necrosed, with a few loose acid-fast germs. 
Around this was a considerable number of lymphocytes, enclosing numerous bacilli, 
and at the periphery organised connective tissue with a few cells filled with bacilli. 
The authors considered that a proliferation of the original germs had taken place. 

1909. Sugai inoculated Japanese dancing mice intraperitoneally with an 
emulsion of fresh leprous nodules, and found as a result the development of miliary 
granulomata on the peritoneal lining, especially in the hepatic region and epiploon. 
Also the bronchial and peritoneal glands showed characteristic leprous lesions. All 
contained acid-fast germs. The attempt to transmit the disease from these to 
other mice failed. 

1909. Kitasato appears to have successfully inoculated an orang-outang on 
the cornea. Only scanty details were, however, given. 

1909. Stanziale reports upon experiments on rabbits. In one typical case 
he inoculated into the eye of the animal a fragment of leproma. During the first 
few days a marked hyperaemia developed round the opening in the cornea. Soon 
afterwards a constant diminution in the size of the nodule could be made out, 
accompanied by the formation of an opalescent exudation, which took its departure 
from the inoculated piece of tissue and spread out as a fan-shaped segment. After 
twenty days the nodule became stationary, and remained so during fifteen to 
twenty days without any noticeable change taking place. After forty days the 
nodule began slowly to increase till it had trebled its original size. In the same time 
small grey nodules made their appearance, and could clearly be distinguished on the 
surface of the inoculated tissue. These little nodules attained the size of a pin’s 
head. Some appeared to be adherent, while others were loose and independent. 
After the initial hyperaemia due to the operation had disappeared, a new formation 
of blood vessels took place, which spun a delicate network round the piece of tissue 
inoculated. After seventy days the rabbit’s eye was enucleated, part of the tissue 
was used for histological observations and the rest was grafted into the cornea of 
two other rabbits. In one the graft got resorbed without any further results in 
the course of two months. In the other rabbit the graft began to increase in size 
after fifty days, and produced similar changes to those described in the foregoing 
animal. The experiments are being continued by the author, but to date the 
results obtained are : thirty-one rabbits have been inoculated, and in eight it was 
possible to produce leprous ocular lesions. In some of the positive cases the 



3 + 


material was taken eighteen to nineteen hours after excision from the patient. In 
all cases in which the inoculation succeeded the Wassermann reaction was positive. 
As a control Stanziale inoculated lepra nodules into the abdomen of another rabbit, 
and found at intervals after a month the reaction constantly negative. The same 
negative result was achieved after grafting into the cornea of rabbits pieces of human 
slrin or dead leprous nodules. 

These experiments of Stanziale have been most carefully carried out, and full 
descriptive details are given ; the only defect is, however, that no metastases in the 
inner organs are mentioned. 

1911. Puval inoculated a series of animals—namely, four rats, four white 
mice, and four Japanese dancing mice—with grumous material taken from an 
acute case of human leprosy which had developed numerous soft subcutaneous 
leprous masses and had repeated attacks of leprous fever. A small quantity of 
material was used in each case—namely 0*5 c.cm. emulsified in 1 c.cm. of normal 
saline solution. Some were injected intraperitoneally, others subcutaneously. 

The two white mice which received intraperitoneal injections died fourteen 
days after the inoculation. At the autopsy both showed a general infection of the 
peritoneum with a pure growth of the M. lepra, while the mesentery, omentum, 
visceral and parietal peritoneum contained numerous minute, firm greyish-white 
nodules, which on microscopic section proved to be typical leprous lesions. The 
most surprising feature was the occurrence of a slightly turbid, semigelatinous 
exudate, which microscopically consisted almost entirely of large mononucleated 
cells (macrophages). Great numbers of these cells were filled with acid-fast bacilli, 
and scarcely any were found that did not contain a few. No remarks are made 
about the other animals. 

1911. Couret fed two goldfish on teased nodules from a leper. No other 
food was allowed the animals until they had disposed of the leprosy tissue, after 
which they were cleansed and transferred to a clean aquarium. 

The first was examined twenty-four days later; acid-fast organisms had not 
been found in the faeces for three days before the animal was killed. No evidence 
of infection was apparent externally or internally. Films made from the omentum 
and other organs showed a few scattered acid-fast bacilli. The average number of 
bacilli to the slide was about four, and no change in morphology was seen. The 
second fish was examined after thirty-seven days. The findings were similar to 
those in the first one. With reference to these experiments, it should not be 
forgotten that tap water appears to harbour acid-fast germs frequently, as shown 
by Brehm, Beitzke, Schern, and Dold. The tap water at the Lister Institute in 
London and of the Public Health Laboratory in Cape Town, also contains acid-fast 
germs; I have also seen them in the taps at Stockholm. (See Plate IV, fig. 20.) 

Serra (1911 and 1913) describes a series of experiments he has carried out on 
rabbits by means of the intra-ocular method. 

His results on the whole seem to correspond to those of Stanziale, but it is 
extremely difficult to unravel any definite statements as to the facts observed 
from a mass of secondary detail. Too much stress seems to be laid on the Wassermann 
reaction observed in some of the rabbits inoculated, and no details are given as to 
way in which it has been carried out; whilst also in this case no metastases in the 
inner organs are mentioned. 

It is to be hoped that other following publications will bring more light on 
the subject. 

Ch. Nicolle and Blaizot succeeded in producing in Macacus sinieus , by inocu¬ 
lating fragments of leprous tissue, local lesions very similar to those of dermal 



35 


leprosy in man. They repeated these experiments on three monkeys (bonnet 
chinois) and one chimpanzee, and reported on the results in 1911. 

One monkey received sixteen inoculations in toto> but none of them ever 
generalised, and the resulting nodules got slowly resorbed in the course of time. 

A second monkey died without showing any visceral lesions. The experi¬ 
mental lepromas were in course of development, and showed numerous acid-fast 
micro-organisms under the microscope. 

A third monkey received nine inoculations at intervals varying between a week 
and twenty-five days. The resulting nodules softened, caseated and broke down. 
The pus showed a few degenerated “ bacilli ” among numerous polynuclear cells. 

The chimpanzee received eight inoculations in the course of four months. 
One injection was followed by the development of a local nodule, which was 
followed later on by several little nodosities, which soon became confluent. This 
lesion only lasted about a fortnight and then got resorbed. 

In these cases, therefore, we had a temporary or negative result in most cases, 
and a localised success in one experiment. 

Bayon (1912) injected four rats on two occasions in the testes with ground-up 
nodules from a case of leprosy from Mauritius. Two rats did not show any macro¬ 
scopic lesions, even after four months. One rat, however, developed, four weeks 
afterwards, a nodule at the site of inoculation, which grew to the size of a small pea. 
On puncture it showed acid-fast germs and necrosed tissue, some of which, no 
doubt, represented the original cells injected. After five weeks the rat died, but 
no acid-fast micro-organisms were found in the organs. Microscopical sections 
were made of liver, spleen, and testes. The disease, if transmitted, was localised. 
The whole of the nodule was then injected subcutaneously into two other young 
rats. Three months after inoculation, one of the rats was losing fur on the surface 
of its abdomen, without there being any signs of the usual rat scabies about the 
ears or tail. Small shotty nodules could be felt under the skin. 

The other rat developed, three months after the second injection, a nodule 
in the left testis. This broke down, and was found to contain acid-fast rods in 
great quantity. This rat was killed four months after inoculation, the slight 
wound caused by the nodule breaking down having healed. At the site of injection 
and in the corresponding inguinal glands numerous acid-fast rods were found. 
One of these small glands was inoculated into the testes of another rat, and on 
killing it six months afterwards no acid-fast micro-organisms were found in the 
testes, but definite deposits were present in the inguinal glands, spleen and liver. 
A multiplication must have taken place, because the acid-fast bacteria detected in 
these organs were evidently more numerous that the relatively small quantity 
injected. (Plate III, Fig. 14.) 

At Robben Island, during 1912, 1913 and 1914, thirty experiments on rabbits 
were undertaken by means of the intra-ocular method. Exact data will be 
published in due course, when the histological study of the resulting specimens is 
undertaken. 

It is, however, worthy of note that one single rabbit of the thirty inoculated 
has shown permanent lesions—that is, minute white, vascularised nodules in 
the cornea and on the iris. This animal was operated upon 1st November, 1912 ; 
nodule from Pat. No. 1294. 

As a rule the remaining animals developed inflammatory appearances similar 
to those described by Stanziale, but in the course of time the inoculated fragment 
of leprotic tissue got resorbed, all signs of inflammation or production of new 
tissue subsided, and in conclusion only a linear scar remained as a trace of the 
operation. 



3 6 

One animal, however, still shows numerous minute leprotic lesions one year 
and seven months after inoculation. 

I do not wish to admit a definite communication of leprosy to these animals, 
till I have been able to achieve the results accomplished with the rats—that is, 
the transmission of the virus through several generations, and deposits in the inner 
organs. 

Reenstierna (1912) injected a Macacus rhesus under the mucous membrane of 
the nose with 0*5 c.cm. of an emulsion made from a leproma containing numerous 
“ bacilli.” The animal died forty-two days after inoculation. The inner organs 
showed no alteration, but the nose had developed a perforation of the septum ; 
moreover, there was extensive ulceration of the upper lip and nasal mucous mem¬ 
brane. The microscope revealed numerous extra-cellularly situated acid-fast rods 
in the nasal secretion and sections of the diseased tissues. In addition an acid- 
labile but Gram-positive filamentary germ was present with fragments which, 
morphologically, were similar to lepra “ bacilli.” 

Another M. rhesus was injected intra-peritoneally and subcutaneously with 
the same emulsion as in the previous experiment. The monkey died fifty-nine 
days afterwards, and showed numerous caseating nodules in the liver, spleen, 
lung and mediastinal lymph-glands. These lesions contained numerous acid-fast 
rods. 

Five guinea-pigs were inoculated with the caseating nodules from the spleen ; 
they died within three to five weeks and showed localised lesions which contained 
very numerous acid-fast rods. 

A third M. rhesus monkey was injected with the same emulsion intraperi- 
toneally and subcutaneously. It died six months afterwards, and developed small 
nodules at the sites of injection which contained acid-fast rods. A cherry-sized 
nodule which developed on the upper-lip contained numerous acid-fast intra¬ 
cellular “ bacilli ” and vacuolated cells. Acid-fasts were also found in the liver, 
spleen and bone-marrow. 

These are the successful or reputed successful attempts to 
transmit leprosy to animals by means of injection or inoculation of 
leprous tissues. 

As will be seen, they are certainly more numerous and have been 
more carefully observed, than the isolated statements in connection 
with the communication of syphilis to animals, which were to be 
found in literature only so short a time ago as the first year of this 
century. And yet syphilis can be communicated to rabbits with 
relative ease. In dealing with leprosy, where even direct inocula¬ 
tion of human beings has failed (except in Aming's case) one can 
but expect to find much more serious difficulties, such as: 

(1) Lack of knowledge in relation to the bionomics of Hansen's 
‘ bacillus.' 

It does not seem to be understood that though attendants in 
leper asylums, and persons living with lepers, must often come in 
contact with numerous ‘ bacilli ’ which we know nodular lepers cast 



37 


about them, yet only a very small proportion ever develop the 
disease in a clinically recognisable form. In other words, in the 
great majority of cases either the ‘ bacilli ’ remain absolutely 
quiescent in the body or get harmlessly eliminated. 

Therefore it should not astonish us that a great proportion of 
the animals inoculated eliminate the acid-fast bacteria injected after 
a longer or shorter period. 

(2) The difficulty of proving whether any ‘ bacilli ’ seen in 
experimental lesions are simple deposits of dead acid-fast bacteria 
or whether they are alive and in course of multiplication. 

This is really an outcome of No. i, as it depends on an 
incapacity in comprehending how extremely passive, inert and 
resistant Hansen’s ‘ bacillus ’ is. 

It is true that Campana, Wesener and others have shown that 
it is possible to bring about lesions in animals very similar to those 
described by the experimenters who claim having transmitted 
leprosy to rabbits or rats, by simply injecting tissues containing 
numerous ‘ bacilli ’ which had been autoclaved or kept in alcohol 
for months. As we know of no method of cultivating directly the 
acid-fast germs seen in leprosy with their original morphology and 
staining properties, it is of course difficult to refute this argument 
except by pointing to the analogy with rat-leprosy, where, though 
we know that the ‘ virus ’ used in experiments is living because it 
•can be indefinitely transmitted, still, except for the latter particular, 
the lesions caused by killed and living virus are extremely similar. 

(3) The similarity existing between the lesions produced in 
rabbits and those found in experimental tuberculosis. 

This is the argument which settled the results of Melcher and 
Ortmann, Wesener and others. At the time leprosy and tuber¬ 
culosis were considered to be as easily distinguishable histologically 
and bacterioscopically as they are clinically. Further study has, 
however, shown that this is not the case, and that under certain 
conditions only experiments with guinea-pigs can decide whether 
we are dealing with lesions brought about by Koch’s or Hansen’s 
' bacillus.’ 

Taking into careful consideration the previous experiments and 
personal experience, in addition to the interpretation of the clinical 
notes made on lepers who have been under observation for fifteen 



3 « 


and more years in South Africa/ no other conclusion can be arrived 
at than that leprosy is directly, experimentally, communicable 
from human beings to rats and rabbits; that a long time of 
incubation must be allowed for; that the great majority of animals 
eliminate the 1 bacilli * injected after a shorter or longer period of 
quiescent deposit. 

We have, therefore, under experimental conditions an analogous 
counterpart of what is happening daily with people who contract 
leprosy, for even prolonged exposure to infection causes the disease 
to develop only in a small percentage. Those who show clinical 
symptoms of the contagion appear to eliminate the majority or 
practically all their 1 bacilli * if they live long enough to become 
what is called an 4 arrested * case. Such an 1 arrested * case is a 
leper who is known to have had numerous visible lesions teeming 
with acid-fast rods, but who for the time being only shows scars 
and traces of the disease with no 4 bacilli.* Such cases relapse at 
times, or may show small bacillary deposits in their inner organs 
at death, but even then there can be no doubt that they have 
succeeded in eliminating or destroying millions and millions of 
4 bacilli * from or in their nodules. 

A somewhat analogous observation is often made at autopsies 
in connection with tuberculosis of the lungs. It is then seen that 
very many bacillary infections of this organ do not develop into 
clinical tubercles, but heal with scarring. 

All these considerations should, and do, give a satisfactory 
explanation of many negative results and condone the scepticism 
with which many successful experiments have been received. 
Conclusive histological specimens like those shown in Plate III, 
fig. 14 cannot be ignored, especially as experimental results agree 
in an exceptional manner with clinical observation. 

Once more, these conclusions have to be arrived at : 

(1) Leprosy is experimentally transmissible to animals such as 
the rabbit and the rat. Allowance must be made for the fact that 
the great majority of inoculations do not succeed, owing to the 
4 bacilli * being eliminated or destroyed without leaving any visible 
trace. 

• This shows that in single instances lepers get rid of nearly all the acid-fast germs from their 
lesions. 



39 


(2) The incubation period is very long, and the resulting 
lesions in the inner organs may not be visible to the naked eye. 

(3) In rabbits the bacillary deposits in the inner organs 
resemble tuberculosis. In such cases the question can only be 
decided by means of guinea-pig inoculations. 


REFERENCES 

(Chronologically arranged) 


Neisser (1881). VVeiterc Bcitrage zur Aetiologie der Lepra. Virchow’s Archiv, LXXXIV, 314. 
Damsch (1883). Ubcrtragungsversuche von Lepra auf Thiere. Virchow’s Archiv, XCII. 

Melcher und Ortmann (1885). Ubertragung von Lepra auf Kaninchen. Berliner klinische 
Wochenschrift, p. 293, No. 13. (1886) Experimentelle Darm und Lymphdriisen Lepra 

bei Kaninchen. ibidem No. 9. 

Wesenek (1888). Zur Frage der Lepraiibertragung auf Tiere. Centralblatt fiir Bakteriologie, 
III, 482. 

Vossius (1888). Zeitschrift fiir vergleichende Augenheilkunde, VI, 1. 

Wnoukow (1893). Inaug. Thesis Kasan (Russian), Ref. Centralblatt fur Bakteriologie, 

XII, 173* 

Tedeschi (1893). Centralblatt fiir Bakteriologie, XIV, 113. 

Iwanow (1902). Sur lc sort des bacilles de la lepre dans l’organisme des animaux (cobayes). 
Annales de l’lnstitut Pasteur, No. 10. 

Y’ amada, Toyama, Kurita (1903). Japanese Journal of Dermatology, III, 529, quoted by 
Sugai. 

Thiroux (1905). Quelqucs tentatives d’inoculation de la lipre. Annals d’hygiene et mid. 
coloniale, VIII, 148. 

Nicolle (1906). Recherches experimentales sur la lipre. Annales de l’lnstitut Pasteur XX, 
389. 

Marchoux et BoURRtT (1909). Recherches sur la transmission de la lipre. Annales de 
l’lnstitut Pasteur, XXXIII, 513. 

Sugai (1909). Nachtrag zur gelungenen Ubertragungsversuchen mit Lepra bei Saugetieren. 
Lepra, VIII, 203. 

Kitasato (1909). Die Lepra in Japan. Zeitschrift fiir Hygiene, LXIII, 507. 

Stan zi ale (1910). Inoculazioni di materiale leproso nella camera anteriore dei conigli. 
Giornale italiano delle malattie veneree, V, 1. 

Couret (1911). The behaviour of bacillus leprae in cold blooded animals. Journal of 
experimental medicine, VIII, 576. 

Serra (1911). Inoculation de culture du bacille de Hansen dans oeil du lapin. Lepra, XII, 1. 

Duval and Gurd (1911). Studies on the Biology of and Immunity against the Bacillus of 
leprosy. Archives of Int. Medicine, VII, 231. 

Nicolle, Ch. et Blaizot (1911). Essai de reproduction de la lipre chez le Chimpanz6 et les 
Singes inferieurs. Archives de l'lnst. Pasteur de Tunis, 275. 

Bayov (1912)- On the transmission of leprosy to animals by direct inoculation. British 
Medical Journal, February 24th. 

Nakano (1912). Uber die kunstlichc Zuchtung von Leprabazillcn in Tierleichen. Archiv. 
fur Dermatologie, CXI, 819. 

- (1912a). Experimentelle Untersuchungen iiber die Infektionsmoglichkeit von 

Japanischen Hausratten, Kaninchen und Meerschweinchen mit Lepramaterial. Ibidem, 
CXIII, 787. 



4 ° 

On Rat Leprosy and its Relation to the Human Disease 

A disease extremely similar to leprosy, not only in its clinical 
features but also from an histological and bacteriological point of 
view, was discovered in rats by Stephansky in Odessa, in 1903. Its 
presence among the sewer rats of Berlin was confirmed by Lydia 
Rabinowitch the same year. Both these observers considered that 
rat leprosy could not be transmitted experimentally from one 
animal to the other, and that the acid-fast micro-organism found 
in enormous quantities in the lesions of the animals affected could 
not be cultivated on nutrient media. 

George Dean had studied this same disease at Elstree, near 
London, for some time, and published his results nearly con¬ 
temporaneously with Stephansky and Rabinowitch. He was, 
however, successful in transmitting rat leprosy through several 
generations of rodents, a result which has since been amply 
confirmed by Marchoux and others. G. Dean’s cultural attempts 
only succeeded in isolating a diphtheroid, which showed filamentary 
forms, and much to his astonishment reacted specifically with the 
serum of lepers. The results of his agglutination experiments 
were : 

1 1. The serum of rats inoculated with rat-leprosy, i.e., with acid-fast 
“ bacilli,” agglutinated the diphtheroid. Normal rat’s serum had no agglutinating 
properties for this micro-organism. 

2. The serum from a case of human leprosy agglutinated the acid-fast micro¬ 
organism from the rat. Normal human serum had no agglutinating power. 

3. Normal human serum and the serum from a tuberculous patient failed 
to agglutinate the diphtheroid, whereas the serum from a case of leprosy had 
distinct agglutinating properties.’ 

These carefully planned tests, in addition to the histological 
appearances, the cultural and experimental behaviour of the micro¬ 
organism causing the disease in rats, lead us to believe that rat 
leprosy is possibly related to human leprosy in a manner similar to 
that existing between bovine and human tuberculosis. 

Marchoux has carried out very carefully-planned experiments 
with the strain of rat leprosy he discovered in Paris, and has come 
to the following conclusions: — 

* 1. That only six in a thousand sewer rats in Paris present marked leprotic 
lesions, but that 5 per cent, harbour acid-fast micro-organisms (bacille de 
Stephansky). 

2. That the inguinal glands appear to be the first attacked by the disease. 



4 1 


3. That the lung acts as a filter in arresting the germs and then directing 
them towards the mediastinal glands. 

4. That by simply placing the virus of the disease on the scarified or depilated 
skin, an infection can be brought about in rats even more easily than by subcutaneous 
injections. 

5. That intact skin or even the bare skin of very young rats does not allow 
the penetration of the virus. 

6. Rat leprosy is a disease peculiar to the rat. Mice can be infected, it is 
true, but with less ease than is the case with rats. Attempts to infect through 
the scarified or depilated skin do not succeed as often as subcutaneous injection in 
mice. In such cases one finds amongst the “ bacilli ” involution forms similar to 
those met with in rats injected with human leprosy. 

7. Mycobacterium leprae murium , like Hansen’s “ bacillus ” is a parasite of 
the mesodermal cells. It does not live at the expense of its host, but of the same 
substances which nourish the cell containing it. 

8. Granular bacilli are dead. 

9. A first culture is relatively easy to obtain. Further sub-plants are not 
obtainable. 

10. The bacillus disappears rapidly in contaminated media (en milieu impur). 

11. It does not resist desiccation. 

12. It is not killed by a temperature of 6o°C. during five minutes. It dies 
after being exposed at the same temperature for fifteen minutes. 

13. Infection takes place through the skin, and accordingly superficial surface 
regions are generally more intensely affected. 

14. Infection follows the lymphatic stream. 

15. The point of inoculation is not always the most heavily infected spot. 

16. Male rats can be infected by depositing some of the virus under the 
prepuce without causing any lesions of the mucous membranes. 

17. However, the spontaneous disease does not seem to be transmitted in 
this fashion. 

18. Insects do not carry the disease. 

19. The sarcoptes of rat scabies play only an indirect role in the diffusion 
of the disease. 

20. The immediate contact of a skin abrasion with a diseased surface or 
with freshly contaminated objects is the usual mode of contagion. 

21. By introducing a large number of micro-organisms through the digestive 
tract a primary pulmonary infection can be produced. 

22. The lymphatic glands of artificially inoculated rats are generally very 
small; those of spontaneously diseased rats, on the contrary, generally much 
enlarged. 

23. Inoculation generally produces an infection of the lymphatic glands. 
Impure virus has to be injected to produce the muscular and cutaneous type of 
the disease.’ 


I have quoted extensively tiiese results of Marchoux and Sofel, 
because not only all experiments have been carefully and 
systematically worked out, but the majority of the observations 
made seem to agree so completely with what we know of the 
pathology of human leprosy. 



4 2 


Currie and Hollmann also carried out experiments with rat 
leprosy, and in two papers on the subject came to the following 
conclusions: 

* i. In some cases of artificially acquired rat leprosy the onset is with broncho¬ 
pneumonia, accompanied by a septicaemia and without any other demonstrable 
lesions. 

2. In other cases of this disease pneumonia is a very early lesion, but we 
cannot positively state that it is always the first lesion. 

3. That the animal may die in the pneumonic stage before other lesions 
present themselves, or it may develop pneumonic symptoms and recover from the 
same only to develop later the well-known lesions. 

4. That during the stage of the disease in which the animal is very ill certain 
mites (Latlafs echidninus) were found to be very numerous on the animals’ bodies. 

5. That during the stage of the disease in which septicaemia is marked, these 
mites* digestive tracts contain the bacilli of rat leprosy in considerable numbers, 
and, therefore, these parasites may be one means of transmitting the disease. This 
latter probability is, of course, not proven.’ 

In a later paper, published in 1912, these authors add the 
following to the previous conclusions: 

‘ 1. In the disease we are dealing with, whether the animal is inoculated by 
a laboratory method or simply allowed to develop the disease from coming into 
contact with infected rats, i.e., the natural mode of infection—the lesions met with 
are practically the same. 

2. With the exception of the local lesion, occasionally produced at the site 
of artificial inoculation, infection of the viscera seems to usually precede the lesions 
of the skin. 

3. Of the visceral lesions, a broncho-pneumonia is often the earliest and 
most constant. Infection of the spleen is also often an early event. 

4. The heart blood of infected rats often contains the bacilli of rat leprosy, 
and no difficulty is experienced in demonstrating the presence of acid-fast bacilli 
in the mites contained on the bodies of these animals when the latter’s heart blood 
contains the organisms. 

5. The fact that these mites contain the bacilli so frequently naturally leads 
one to suspect that they may be one of the means of transmitting the disease from 
rat to rat, but up to the present time we have no evidence that such is the case. 

Attempts to Cultivate the Bacillus of Rat Leprosy 

As reported in our first paper, we have made many attempts to cultivate the 
bacillus of rat leprosy on artificial culture media, both on the media ordinarily 
employed in the laboratory and by the method of Clegg in symbiosis with amoeba 
and cholera. We have now to report that we have continued these experiments 
for‘nearly a year, and that our results have been entirely negative, not having 
secured a single culture of an organism which we considered to be the bacillus of 
rat leprosy. In one of our attempts to cultivate this organism from the ulcer on 
the abdomen of a leprous rat, we succeeded in growing what appeared to be an 
acid-fast streptothrix. As one might expect to obtain such organism on an 
exposed surface of that kind, we regarded this culture as an accidental contamina¬ 
tion, and not the bacillus of rat leprosy.’ 



43 


It is hardly feasible to bring these results in correlation with 
anything observed in human leprosy, except for the fact that also 
these authors have found the disease can be transmitted by means 
of inoculation from one animal to another, an experiment which 
did not succeed in the hands of earlier observers. In a similar 
fashion the transmission of human leprosy to animals has failed 
time after time, and yet under exceptionally favourable circum¬ 
stances it can be successfully achieved. The repeated attempts to 
cultivate the micro-organism of the disease, which only resulted in 
the isolation of acid-fast filamentary, branching bacteria (diph¬ 
theroids are not mentioned) which were considered to be 
contaminations, are also worth noting, because this observation 
coincides with the results of much of the work done on the 
bacteriology of human leprosy. 

Bay on (1912) succeeded in isolating on fish-juice-agar from the 
spleen of a rat which had been injected with a ground-up nodule 
from a spontaneously diseased rodent, a moist, ivory-white, creamy 
growth, which multiplied in sub-cultures very slowly at 37 0 C. on 
the media generally used for the artificial cultivation of tubercle. 
This culture produced in rats the identical glandular and visceral 
lesions observed in rats injected with the virus from spontaneously 
diseased animals. The glands and organs infected were teeming 
with acid-fast micro-organisms, which caused little or no tissue 
reaction and a very minimal production of giant cells (Plate III, 
figs. 16 and 17). 

The culture does not grow at room temperature or on common 
gelatine.' It is difficult to isolate again from the inner organs of 
the animals into which it has been injected. Guinea-pigs, fowls and 
rabbits do not show any lesions if inoculated with small quantities 
(five to ten loops), but if a culture slope or more is introduced intra- 
abdominally, after a time caseating, necrosed nodules are produced 
on the surface of the peritoneum and the omental glands can be 
considerably enlarged, showing big masses of acid-fast micro¬ 
organisms. The necrosis and caseation are partially due, I believe, 
to the mechanical foreign-body action of the large numbers of 
resistant micro-organisms injected. 

No growth could be obtained under anaerobic conditions. 

In an oxygen atmosphere the cultures develop somewhat more 



44 


rapidly; after ten to fourteen days a thick growth is discernible, 
whilst under ordinary conditions, at 37 0 C., three or four weeks are 
needed. 

No multiplication was obtained on gelatine at room temperature 
or on common agar in the incubator. 

No pigment was produced on any of the following media: 
fish-j uice-glycerine-agar, glycerine-agar, glucose-agar, glycerine- 
potato, rice-glycerine-milk, sterilised wedges of rabbit’s liver + 
glycerine, Dorset’s egg-medium. All these media were suitable for 
growing the micro-organism. 

Morphologically the culture is a short rod, with a marked 
tendency to pleomorphism; clubbed shapes are frequently seen, and 
at times true branching can be detected. The length of the rods 
varies; in fact, the microscopical appearances are very similar to 
those of a culture of avian tuberculosis, which it also resembles 
from a cultural point of view. 

It is acid-fast, i.e., cannot be bleached by one minute’s treatment 
with 20 per cent. H 2 S 0 4 after having been stained for five minutes 
with warm carbol-fuchsine. It is also alcohol-fast. After soaking 
for six hours in warm alcohol its staining properties are unimpaired. 

It is Gram positive. 

Very numerous attempts to isolate the same micro-organism from 
several other cases of rat leprosy failed completely. Marchoux is 
therefore not quite correct in saying that: ‘ Bayon, au contraire, 
l’aurait cultiv6 assez facilement.’ (l.c. p. 23.) 

Further cultural results have been published by Hollmann, 
Chapin, Zinsser and Carey, Marchoux and Sorel, and, of course, 
Wellman. None of these cultures have succeeded in producing 
leprotic lesions. 

After Stephansky’s and Rabinowitch’s and Dean’s observations, 
rat lepra was found in Australia by J. R. Bull, in New South Wales 
by Tidswell, in the United States by Wherry, McCoy, Walker, in 
Roumania by Mezinescu and Alexandrescu, in Japan by Jitoyo and 
Sakaki, in Paris by Marchoux, in New Caledonia by Leboeuf, in 
Ipswich by Petrie and Macalister (see PI. I, fig. 4), in India by the 
Plague Commission, in Queensland by Priestley. 

Though rat lepra seems to be spread nearly all over the world, 
still Brinckerhoff examined 16,000 rats in the Hawaii Islands 
without finding any trace of an animal affected with the disease. 



45 


In Cape Town I have had now 1,378 sewer rats examined 
without having succeeded in a single instance in discovering a 
marked case of the disease.* 

Dr. G. W. Robertson remembers, however, to have noticed one 
animal the lesions of which contained numerous acid-fast rods, in 
the course of plague investigations, when about 30,000 rats were 
examined. 

Mezinescu succeeded in preparing from the ground-up nodules 
of leprous rats an antigen which was capable of absorbing comple¬ 
ment when linked up with the serum of lepers; a further proof of 
the relationship of both diseases, though, on the other hand, it 
must be admitted that the serum of lepers is similar to that of 
syphilitics in this respect, because it is capable of reacting with 
specific and certain non-specific antigens. A distinction can only 
be made by carefully titrating all the elements of the haemolytic 
system, and varying the proportions of antigen and antibody in a 
suitable fashion; up to the present this has only been done in quite 
a minimal proportion of cases. 

Rats are not the only animals affected by a disease akin to 
leprosy. Acid-fast micro-organisms which belong to the tubercle 
group, but are not pathogenic for guinea-pigs in small quantities, 
and are only cultivable with extreme difficulty, cause diseases in 
cattle, horses, sheep, and possibly goats. The cattle disease has 
been named by K. F. Meyer 4 Enteritis hypertrophica bovis 
specifica.* 

Twort and Ingram, Vukovic, MTadyean, McGowan, have 
described 1 Johne’s disease* in the sheep. This disease differs from 
rat-leprosy in so far that the symptoms are mostly confined to the 
digestive tract. 

REFERENCES 

(Chronologically arranged.) 

Stephansky, W. K. (1903). Eine lepraahnliche Erkrankung der Haut und dcr Lymphdriisen 
bei Wanderratten. Centralblatt fur Bakteriologie, XXXIII, 481. 

Rabinowitch, L. (1903). Uber eine durch saure-feite Bakterien hervorgerufene Haut- 
erkrankung der Ratten. Centralblatt fiir Bakteriologie, XXXIII, 577. 

Dean, G. (1903). A disease of the rat caused by an acid-fast Bacillus. Centralblatt fiir 
Bakteriologie, XXXIV, 222. 

- (1905). Further observations on a leprosy-like disease of the rat. Journal of Hygiene, 

v, 99- _ 

*My thanks are due to Dr. Mitchell and Dr. J. Anderson of Capetown for procuring 

these rats. 




4 6 


Tidswell, F. (1904)* Note on a leprosy-like disease in rats. Report of the Board of Health 
on Leprosy in New South Wales, p. 13. 

Reports on Plague Investigation, etc. (1907). Journal of Hygiene VII, 337. 

Bull, J. R. (1907). Leprosy-like disease of the rat. Intercolonial Med. Journal of Australasia, 
May 20th, p. 215. 

Wherry, W. B. (1908). The leprosy-like disease among rats on the Pacific Coast. Journal of 
the American Medical Association, L, 1903. 

- (1908a). Notes on rat leprosy and on the fate of human and rat lepra bacilli in flies. 

Journal of Infectious Diseases, V, 307. 

- (1909). Experiments on vaccination against rat leprosy. Ibidem, VI, 630. 

McCoy, G. W. (1908). Leprosy-like disease in rats, U.S.P.H. and Marine Hospital Service 
Reports, July 10th. 

- (1908a). Distribution of the leprosy-like disease of rats in San Francisco. Ibidem, 

November 6th. 

- ( I 9 I 3 )* Observations on naturally acquired rat-leprosy. Ibidem, No. 61, XXIV, 

27-30. 

Mezinescu, D. (1908). Maladie 16 preuse des rats et ses relations avec la lipre humaine. 
Comptes rendus de la Soci6t6 de Biologie, LXIV, 314. 

- (1909). Ibidem, LXVI, 56. 

Walker, A. (1918). A report on some cases of rat leprosy. Journal of the American Medical 
Association, LI, 1153. 

Alexandrescu, J. (1908). Lepra sobolanilor. Quoted from Marchoux (1912). 

Kitasato (1909). Die lepra in Japan. Zeitschrift fur Hygiene, LIII, 507. 

Brinckerhoff, W. R. (1910). Rat leprosy. U.S. Public Health Bulletin, No. 30. 

Currie, D. H. and Hollmann, H. T. (1910). A contribution to the study of rat leprosy. 

U. S. Public Health Bulletin, No. 41. 

- (1912). XVII. Further observation on rat leprosy. Ibidem, No. 50. 

Jitoyo, K., and Sakaki, R. (1910). On rat leprosy. Ref. Centralblatt fur Bakteriologie, 
LVIII, 690. 

Ehlers, Bo urret et With (i 91 i). Recherches sur le mode de propagation et les proc£des de 
diagnostique bact6riologique de le lipre. Bulletin dc la Soci6t6 de Pathologie Exotique. 

3 *» 2 53 - 

Leboeuf, A. (1912). Existence de lepra murium en Nouvelle Caledonie. Bulletin de la 
Soci6t6 de Pathologie Exotique, No. 7. 

Marchoux, E., et Sorel, F. (1912). La Wpre des rats. (Lepra murium.) Annales de 
l’lnstitut Pasteur, Aout. 

-(1912a). Lepra murium. Comptes rendus de la Society de Biologie, LXXII, 

159, 214, 269. 

Marchoux, E. (1912). Role des infections secondaires dans le developpement de la lipre du 
rat. Bulletin de la Society de Pathologie Exotique, p. 466. 

- (1912a). Human leprosy and rat leprosy ; a discussion of their respective problems. 

Trans. Soc. of Trop. Medicine, V, 184. 

Bayon, H. (1912). The culture and identification of the germ of leprosy and the relationship 
of the human disease to rat leprosy. Transactions of the Society of Tropical Medicine, 

V, 158-167. 

-(1912a). A comparative experimental study of the cultures of Clegg, Duval, • 

Kedrowsky, Rost and Williams. British Medical Journal, Nov. 2nd. 

Meyer, K. F. (1912). Bacillus para-tuberculosis Johne. Proceedings Philadelphia Pathol. 
Society, October. 

M’Gowan (1914). A case of Johne’s disease in the sheep, in ‘Monograph on Scrapie,* page 
113-U4. 

Wolbach and Honeij (1914). A critical review of the bacteriology of human and rat-leprosy. 
Joum. of Med. Research XXIV, pp. 367-423. 



+7 


VII. THE SEROLOGY OF LEPROSY AS A MEANS OF DIAGNOSIS 
OF THE DISEASE AND FOR THE PURPOSE OF CONTROLLING 
SPECIFIC TREATMENT 

Soon after the adaptation of Bordet-Gengou’s complement 
fixation to the diagnosis of syphilis by Wassermann and his 
co-workers, Eitner modified this test to suit the special conditions 
of leprosy, and prepared his antigen by means of a saline extract 
from a leproma. He found that the blood of lepers contained, as 
he thought at the time, specific anti-bodies, and accordingly was 
able to bring about the absorption of complement when linked up 
with an antigen prepared according to his method. In the course 
of time it became evident that Wassermann’s test for syphilis could 
not claim to be absolutely specific* Accordingly Eitner amplified 
his experiments, and found that the serum of lepers was also 
capable of giving a positive reaction when linked up with the 
alcoholic extract from a guinea-pig’s heart, and that therefore, also 
in this particular instance, the serum of lepers and that of 
syphilitics behaved in a similar fashion. 

Numerous and various experiments were then carried out with 
different antigens by several investigators, and the serum of lepers 
was carefully compared with that of syphilitics and tubercular 
patients. As far as the original Wassermann test of syphilis is 
concerned, its relation to leprosy can be aptly summed up in the 
conclusions of Howard Fox (l.c.): — 

‘ I. A positive Wassermann reaction is frequently obtained in cases of 
leprosy giving no history or symptom whatever of syphilis. 

2. The reaction is at times very strong, inhibition of haemolysis being com¬ 
plete. 

3. The reaction occurs chiefly in the tubercular and mixed forms of the 
disease, especially in advanced and active cases. 

4. In the cases of the maculo-anaesthetic and purely trophic type the reaction 
is generally negative. 

5. The value of the test is not affected in the slightest by the results found 
in leprosy.’ 

On the other hand, if we consider the test being made with an 
antigen prepared in various fashions from the nodules of lepers 
(Eitner’s test or its modifications), one finds on reviewing the 
literature on the subject that, with a few differences, the results 
mainly agree with Jeanselme’s conclusions, which are the following: 



48 

‘ i. Eitner’s reaction applied to syphilitic sera is positive in 85 per cent, of 
all cases examined. 

2. Eitner’s reaction with the serum of healthy individuals or that of syphilitics 
in whom the disease is quiescent: 100 per cent, negative. 

3. Wassermann’s reaction applied to lepers: Practically the same conclusions 
as Howard Fox.’ 

Eliasberg has shown that the serum of lepers is at times capable 
of inhibiting haemolysis by itself, even small quantities as 0*4 c.c. 
being sufficient. A similar observation has been made by Ehlers 
and Bourret. 

Noguchi’s 4 Luetin * reaction was found to be negative in eleven 
lepers who were free from obvious signs of syphilis (but gave a 
Wassermann reaction) by Clegg. 

Incidentally it can here be mentioned that, according to 
Rocamora’s and also Jeanselme’s experience, salvarsan has no 
influence whatever on the Wassermann reaction in lepers. 

Evidently neither Wassermann’s nor Eitner’s test can be 
considered strictly specific for syphilis or leprosy. It remains, 
however, to be seen whether a test could be devised in which by 
using pure cultures of ‘Hansen’s bacillus’ a clear, specific, definite 
reaction might be obtained. 

Currie and Clegg approached the problem with the intention of 
proving the identity of the acid-fast bacteria they had cultivated 
from lepers, and after numerous experiments concluded that: 

4 1. They were unable to differentiate by the method of Bordet and Gengou 
the leprosy bacillus from certain other acid-fast mirco-organisms. 

2. Extracts of certain acid-fast bacilli, other than B. leprae , will deflect the 
complement when combined with leper’s serum. 

3, They were able to produce specific agglutinins for B. leprae by injecting 
a horse with the cultivated lepra bacilli.’ 

The explanation of these results is given by Much and his 
co-workers, Hossli, Lescke and Deilmann, who have shown that all 
acid-fast micro-organisms are more or less closely related and their 
chemical constituents are similar. Much sums up his results in the 
following words (l.c. 1912): 

* We were able to show—chiefly by the reaction of immune bodies—a relation¬ 
ship between tubercle and lepra bacilli and other acid-fast bacilli, which is due to 
inherent qualities common to all these organisms. Thus tuberculous and tubercle 
immunised persons responded to the reaction of complement-fixation, not only 
with tubercle, but also with other acid-fast bacilli, though not equally with all, 



49 


and with certain sorts more than with others. A table which I have prepared 
shows the quantitative differences in these reactions, and according to this table 
the lepra bacilli have the closest resemblance to the tubercle bacilli. 

In the same way leprous sera react not only against lepra bacilli, which we 
obtained from leprous tubercles, but they give also a specific reaction in the same 
scale of relationship with tubercle and (other) non-pathogenic acid-fast bacilli. 
These reactions are absolutely specific.’ 

Much’s observations and experiments show clearly that, as was 
to be expected, the specific result of any serological test carried out 
with acid-fast micro-organisms is apt to be masked by the inter¬ 
ference of a group-reaction. This is similar in its effects to that 
observed on trying to differentiate from a serological standpoint 
various bacteria of the typhoid or para-typhoid group. It is no 
doubt also analogous to the results of precipitation tests when 
being adapted to identify the blood of closely-related animals. 




■'DOOOS^i 

(•)(•) 

• 9 ji 




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Fig. i. 

Diagram showing two complement fixation tests for the identification of acid-fast micro¬ 
organisms isolated from lepers. 

Both serum and antigen are diluted in falling dosis and mixed in varying proportions. 

Clear circles indicate complete haemolysis, dark circles absolute inhibition ; spots of varying 
sizes correspond to the stages between haemolysis and inhibition. 

Dark circles in the middle vertical line are controls of antigen, serum, complement. 

H. R. Dean having shown that this difficulty can be obviated 
by linking up antigen and antibody in varying dilutions, Bayon 
carried out some experiments on the same lines with the intention of 
getting rid of the group-reaction without having recourse to 




Weichardt’s complicated technique. . These tests require the 
greatest care, because the antibodies ot leprous sera are never very 
strong, and dilutions capable of reacting above i /150 are the 
exception. The antigen was prepared by alternately freezing and 
rapidly thawing, and then shaking mechanically, various acid-fast 
cultures in distilled water. The principal factor consisted in the 
variation of the proportions of antigen and antibody, which can be 
best explained by referring to the figure on p. 49, in which the dark 
rings represent complete haemolysis and the clear rings complement- 
absorption. It will be seen that in all the rows the leprous serum 
was diluted in falling quantities, 1/5, 1/10, 1/20, etc. In the first 
row the antigen was used undiluted, in the second 1/2, in the third 
1 /4, and so forth. By this method not only a control was achieved 
unobtainable by any other technique, but, moreover, very satisfac¬ 
tory and reliable results were arrived at. It was found possible to 
distinguish, from a serological point of view, cultures of various 
acid-fast bacteria, unless they were too similar or closely related, 
e.g., bovine and human tubercle. In this fashion Kedrowsky’s 
strain of leprosy was found to react specifically with the scrum of 
lepers in a dilution of antigen which gave no result with Duval’s or 
Rost’s micro-organisms. 

This test can be applied to the diagnosis of leprosy, but 
unfortunately, in common with many other laboratory methods, it 
fails in those cases of early anaesthetic leprosy, which are the most 
difficult to diagnose clinically. 

In the instance illustrated in fig. 1 the serum of a nodular leper 
was tried against antigens made with the leprosy cultures of 
Kedrowsky and Duval. It will be seen that though Kedrowsky’s 
micro-organism reacted even in a dilution of 1/4 with serum diluted 
1/40, Duval’s bacterium ceased to give any absorption after mixing 
1/2 with serum 1/5. These results have lately been confirmed by 
Kritschewsky and Bierger, who however did not grade their 
dilutions. In the second series of fig. 1 we see a test carried out 
with the serum of a rabbit which had been repeatedly injected 
intravenously with several slopes of Duval’s culture. The serum of 
this animal was used against an antigen made with Duval’s 4 lepra,’ 
and another consisting of Moeller’s 'mist-bacillus’ isolated from 
horse-dung. The result is that very little difference can be made 



out, showing that Duval’s so-called 1 leprosy-culture ’ is very closely 
related to a harmless, ubiquitous saprophyte. 

These results were confirmed in course by the fact that the 
etiological significance of this isolation had to be abandoned by its 
author. 

Moellers (1913) examined the sera from thirty-two lepers, using 
as antigen the following products: Alttuberkulin Hochst, Perl- 
suchttuberkulin Hochst, Tuberkelbazillenemulsion T.O.I., Neues 
Tuberkulin T.R. He sums up his results in the following fashion: 

(1) The serum of mixed and nodular lepers gave a positive 
fixation of the complement in 95 to 100 per cent, of cases examined 
when tested against various tuberculins. Anaesthetic varieties of 
the disease reacted only in 25 per cent. 

(2) In the serum of lepers the amboceptors for bacillary 
emulsions are more developed than those for preparations made 
from the cultural liquid. 

(3) It is not possible to draw any conclusion as to the presence 
of tuberculosis in a leper who shows the presence of 4 tuberculous 
antibodies’ in his serum. 

(4) The complement absorption properties of leprotic sera 
tested with tubercular substances are in direct proportion to the 
extent of mixed and nodular features in lepers. Accordingly it 
appears that this inherent quality of the serum depends on the 
greater or lesser spread of the leprotic lesions (Krankheitsherde); 
this is confirmed by the observation that the reaction was negative 
in arrested cases. 

Kritschewsky and Bierger (1913) found that the serum of lepers 
gave a definite positive complement fixation test with an antigen 
made with Kedrowsky’s culture. Duval’s 1 lepra ’ did not react in 
the same remarkable manner. A close similarity in relation to 
serological tests was found to exist between the leprosy * bacillus ’ 
taken from tissues and that isolated by Kedrowsky. M. tuber¬ 
culosis appeared also to be closely related to the latter micro¬ 
organisms. 

These authors conclude that Kedrowsky’s isolation is true 
leprosy, and that Duval’s chromogenic culture is not etiologically 
connected with the disease. 



5 2 


Gaucher and Abrami, Sugai and others, have tested the 
agglutinating properties of lepers* sera towards Hansen’s ‘ bacillus * 
separated from ground-up nodules. Such experiments are some¬ 
what vitiated from the fact that acid-fast rods are prone to show 
spontaneous clumping. 

Summing up these different independent observations it cannot 
be definitely stated that it is not possible, with due care and 
attention, to carry out serological tests showing the etiological 
relationship of any given micro-organism to leprosy. In the case 
of acid-fast bacteria the group-reaction must be obviated. The 
difficulties are superior to those encountered with other germs; they 
are, however, not insurmountable, as Harris and Langford are 
inclined to believe. 


Tuberculin and Similar Tests in Leprosy 

Abraham, Goldschmidt, Babes and Kalindero, observed practi¬ 
cally at the same time, in 1891, that lepers react with a rise of 
temperature to injections with ‘ Alt-tuberkulin.* Though this 
observation was repeatedly confirmed, the tendency was to consider 
that the reaction was due to a co-existent tuberculous infection. 
Babes, however, examined seven lepers after death, who during life 
had reacted to tuberculin, and found in four chronic tuberculosis 
of the lungs, whilst the remaining three were without any signs of 
infection with M. tuberculosis. Guinea-pig experiments were 
negative. 

Sir Malcolm Morris and Colcott Fox have observed very violent 
reactions in anaesthetic lepers; in the former case bullae appeared 
all over the arms. 

Babes, having given considerable attention to the subject, is of 
the opinion that some lepers react with quite a small dose, others 
only after repeated larger injections. A general reaction takes 
place after about twenty-four hours, in some instances, however, 
after eight to ten, or even as short a time as only two hours, and as 
a rule lasts longer than in the case of tuberculosis. The first can 
be followed by a second or even third rise of temperature. Under 
certain circumstances an extremely marked reaction may follow an 



53 


injection of less than one mmg., lasting several weeks and 
endangering the life of the patient. 

Jadassohn has, however, been unable to produce any reaction 
with tuberculin in four cases he examined. It will be seen that the 
evidence is contradictory, but if we take into consideration the 
relative close relationship of leprosy and tuberculosis from many 
points of view, it cannot appear improbable that lepers should react 
to various tuberculin tests. 

In previous communications Bayon was inclined to consider that 
intra-dermal test after Mantoux’s method, carried out with a 
cultural extract made from Kedrowsky’s strain of leprosy, might 
have a specific and diagnostic value. Further investigations have, 
however, shown that similar local and general reactions can be 
brought about by the extracts of several acid-fast micro-organisms 
such as M. phlei y Moellers ‘ mist-bacillus ’ and Duval’s so-called 
‘ lepra.’ It is true that as a rule the reaction is not as marked with 
the latter substances, but the difference is only one of degree. 

On the other hand, lepers who do not react to a tuberculin test 
may show very definite local reactions after intra-dermal injection 
with one c.cm. of a io per cent, dilution of the extract from 
Kedrowsky’s culture. This may consist of a copper-coloured patch 
appearing at the site of injection or in more marked cases in the 
production of blisters and infiltration (Plate V, figs. 3 and 4). 

One patient at Robben Island, under observation of Dr. W. L. 
Stuart, developed reddening and infiltration and a rise of tempera¬ 
ture to ioo° F. even after injection of minimal doses corresponding 
to one c.cm. of a I /250 dilution of cultural extract. Other nodular 
lepers again did not react even after injection of 3 c.cm. Both 
extremes are exceptional; as a rule one or two c.cm. will be 
followed by a rise of temperature and an intra-dermal injection 
will result in varying degrees of discoloration. Pattison injected 
ten lepers (four nerve cases, four mixed and two nodular) with 
one c.cm. 1/10 dilution; all reacted either locally or generally. The 
local reaction consisted usually of a dark patch at the site of 
injection surrounded by a red zone, which might be swollen and 
tender. 

All these tests have, it is true, confirmatory value, but at the 
present stage they are insufficient as a decisive help to diagnosis. 



54 


REFERENCES 


Akxrbzrg, Almxvist und Jundell (1910). Weitere Beobachtungen uber Wassermanns Serum 
Reaktion bei Lepra. Lepra, IX, 79. 

Babes (*909). Sur des reactions reputes comme specifiqucs de la Wpre. Lepra III, 321. 

- (1910). Uber spczifischc Reaktionen bei Lepra. Zcitsch. f. Imro. u. exp. Thcrapie, 

VII, 578. 

Babes ct Busila (1909). L’cxtrait des lcpromes gardes depuis des annccs dans Talcool comme 
antigine lipreux. C.R. Soc. dc Biologic, LVII, 817. 

- (1910). Etude sur le rapports qui existent entre les antigens et les anticorps 

syphilitiques, tuberculeux et lipreux. C.R. Soc. dc Biologie, LVIII, 181. 

Biehler and Eliasberg (1910). Komplementbindung bei Lepra mit leprosem Antigen. 
Lepra, IX, 207. 

Bruck und Gessner (1909). Uber Serumuntersuchungen bei Lepra. Berl. ldin. Wochen- 
schrift, XLVI, 589. 

Currie and Clegg (1912). Immunity. Lepra, XIII, 10-16. 

Duval and Gurd (1911). Studies on the biology of and immunity against the bacillus of 
leprosy. Arch, of Int. Medicine, VII, 230-245. 

Ehlers et Bourret (1909). Reaction de Wasserman dans la lipre. Bull. Soc. de Path. 
Exotique, 520. 

Eitner (1906). Nachweis von Antikorpem im Serum eines Leprakranken mittels K.om pie- 
mentablcnkung. Wiener klin. Wochenschrift, No. 5. 

- (1908). Zur Frage der Anwendung der Komplementsbindungsreaktion auf Lepra. 

Wiener klin. Wochenschrift, XXI, 729. 

Eliasberg (1909). Komplementablenkung bei Lepra mit syphilitischem Antigen. Deutsche 
med. Wochenschrift, XXXV, 1922. 

Fox, H. (1910). The Wasserman and Noguchi Complement-fixation test in Leprosy. Am. 
Journ. of Med. Science, May. Ref. Lepra, 1912, VIII, 107. 


Frugoni (1909). Syphilis und Lepra. Arch. f. Dermat. u. Syph., XCV, 223. 

Frugoni and Pisani (1909). Vielfache Bindungseigenschaften des Komplements einiger Sera 
(Leprakranker) und ihre Bedeutung. Berl. klin. Wochenschrift, XLVI, 1531. 

Gaucher et Abrami (1909). Le scrodiagnostique des formes atypiques de la lfpre. Lepra, 
VIII, 152. 

Jeanselme (1912). Cytologie ct serologic dc la l£prc. La Presse medicale, 27 Juillet. 

Jundell, Almkwist und Sandmann (1908). Syphilisreaktion bei Lepra. Centralbl. f. inn. 
Medizin, No. 48. 

Kleinschmidt (1910). Bindung Komplementbinden der Antikorper durch Fette und Lipoid- 
korper. Berl. klin. Wochenschrift, XLVII, 57. 

Kritschewsky und Bierger (1913). Zur Frage uber das Verhaltnis des Bacillus leprae Hansen 
zu einigen bie Lepra gezuchteten Mikro-organismen. Zeitsch. f. Hygiene, LXXIIT 
509. 


Lewin, A. (1911). Wasserman’s reaction in lepers. Russki Wratch. Ref. Lepra 
XII, 229. 

Meier, C. (1909). Serologische Untersuchungen bei Lepra. Lepra, III, 334. 

MSllers, B. (1913). Serologische Untersuchungen bei Leprosen. Vcroffentl. 
Stiftung, Heft VIII-IX, 122. 

Much (1912). Serological and experimental Studies on Leprosy. Trans. Soc. Ti 

V, « 75 - 

- — (1912a). Uber Fettantikorper und ihre Bedeutung (mit besondercr Be 
der Lepra). Beitrage zur Klinik d. Infekt. u. z. Immunitatsforschuncj 
Kabitzsch. 


ir 

* 

2 *- 


r.'rf 


Jr 


Montesanto, Denis et Sotiriades (1910). La sero-reaction dc Wasserma 
Upre. Presse med., 31 Aout. 





55 


Peyri-Rocamora (1912). Lc Salvamn dans la Wpre ; son influence sur le Wasserman dans 
cette maladie. Lepra, XIII, 4-9. 

Phonites et Michaxlides (1912). La sero-reaction de Wasscrmann et cuti-reaction de Pirquet 
dans la Wpre. Lepra, XII, 207. 

Recio, A. (1909). La reaction de Wassermann dans la Wprc. Sanidad y Beneficicncia Habana. 
Sept. p. 292. Ref. Lepra, 1910, IX, 183. 

Sf.rra (1939). La sicrodiagnosi di Wasserman nella Lepra. II Policlinico, No. 12, XVII. 

Spindler (1912). Bcmerkungen uber.den Komplcmentgchalt und die Wasscrmannsche 
Reaktion des Blutes Leproser. Dermatol. Zcntralblatt, XVI, No. 3. 

Slattneano et Danielopolu (1908). Sur la presence d’anticorps specifiqucs dans le serum de 
malades atteints de la lipre. Comptes Rendus Soc. de Biologie, 309. 

- (1908a). Reaction de fixation dans la lipre en employant la tuberculine comme 

antiginc. Ibidem, LXV. 

- (1908b;. Reaction de fixation avec le serum et le liquide cephalo-rachidien des malades 

atteints de lipre en presence de l’antigcne specifique syphilitique. C.R. Soc. de Biologie, 

347- 

SrcAi (1909). Zur klinisch-diagnostischcn Vcnvertung der Komplementsbimlungsmcthode 
bei Lepra. Archiv. fiir Dcrmat. und Syph., XCV, 313. 

Stepfenhagen (1910). Ubcr Komplementsbindungs reaktion bci Lepra. Bcrl. klin. 
Wochenschrift, XLVII, 1362. 

Thomsen und Bjarnhjedinson (1910). Untcrsuchungcn ubcr Komplcmcntbindung mit 
dem Scrum Aussiitziger. Lepra, IV, 191. 

Weciiselmann und Meier (1908). Wasscrmannsche Reaktion in einem Fallc von Lepra. 
Deutsche med. Wochenschrift, XXXIV. 1341. 


VIII. COMMUNICABILITY AND CONTAGIOUSNESS OF 

LEPROSY 

Exact and repeated observations carried out during the last 
thirty years have definitely proved that leprosy is an infectious or 
communicable disease. The 4 wisdom of centuries/ to quote 
Sambon’s expression, had never doubted this fact. It is only due 
to superficial clinical observers of the XIXth century that hereditary 
and other irrelevant factors have lately been at all considered, even 
for a short time, in connection with the disease. 

The statement is often made that nothing is known about the 
way in which leprosy is communicated from the diseased to the 
healthy; but as a matter of fact, the question has been carefully 
studied and the results are quite as definite, if not more so, than 
any obtained in many other infectious complaints, such as measles 
or scarlet fever. 

Our knowledge of the contagiousness of leprosy is founded on 
the following observations: 

(i) That it is a disease due to a definite specific micro¬ 
organism which has only been found in lepers, their excreta or 
immediate surroundings. 



56 


(2) That in Northern Germany, where leprosy had been 
re-introduced from Russia in modem times, it was seen to have 
spread very slowly and concentrically around the first imported 
cases and their contacts. 

(3) That on the island of Oesel, the population of which had 
practically remained stationary during the last fifty-five years, 
Lohk, Talwik, and Dehio were able to show that out of sixty-three 
cases of leprosy only in eight could previous contact with lepers not 
be traced. 

(4) That with one exception (Da Costa) all cases, in which 
contact cannot be proved, originate in countries or districts where 
leprosy is fairly common; but that in the relatively rare instances 
where leprosy has been contracted in countries in which the disease 
is not indigenous, such as the United Kingdom, Southern Germany, 
Holland, it is possible to prove the intimate contact with other 
lepers, who in their turn had contracted the disease abroad. 

(5) In countries where leprosy is rare, such as the Valais, the 
Riviera, Alpes Maritimes, the disease is always found bound to 
definite foci or families. 

(6) That the countries which have carried out a thorough and 
efficient system of segregation have been rewarded by a gradual 
and constant diminution of the disease. (Norway, Iceland, 
Germany, Sandwich Islands, Philippine Islands.) 

(7) That where segregation has been abandoned, or loosely 
carried out, or not enforced at all, the scourge has usually attacked 
an ever increasing number of individuals. (India, Basutoland, 
Dutch Indies). 

The grounds on which the contagiousness is denied are, I 
believe, the following: 

(1) That medical men and attendants never contract the disease 
in leper-asylums. 

This statement is not quite correct. Infection under these 
circumstances is not frequent, but still several cases can be brought 
forward showing that attendants and medical men do contract the 
disease. In South Africa the Medical Superintendent of a Leper 
Asylum contracted leprosy. Further instances of professional 
infection have been published by Ehlers, Vidal, Jeanselme, 
Nicolas, and others. 



57 


In one case of which I know, a missionary to lepers, trusting no 
doubt to the low degree of infectivity of the disease, allowed his 
little girl to play with leper children. She developed the malady 
whilst at school in Europe. 

Published and unpublished cases of infection among religious 
persons who have attended lepers are also known to exist. (Three 
priests at Molokai, Father Boglioli in New Orleans, two nuns, a 
missionary, a lady missionary.) 

On similar grounds it has been attempted to deny the 
infectiousness of tuberculosis, the deduction being based on 
statistics showing the low ratio of marital and professional 
infection. 

(2) Because the disease cannot be transmitted to animals, and 
inoculation experiments on human beings have failed. 

This also is not quite correct. The first part of this argument 
would have allowed us fifteen years ago to deny the contagiousness 
of syphilis. As a matter of* fact, leprosy is transmissible to 
laboratory animals such as the rabbit or rat, as repeated experi¬ 
ments have shown. That this possibility is not more generally 
recognised is due to the fact that the lesions may be localised and 
fail to develop at all in a very great proportion of experiments, and 
often resembles tuberculosis to a certain extent. 

As to the inoculations which Danielssen, Profeta, Cagnina, 
Bargilli, carried out on themselves and others, the results, it is 
true, were negative, but in view of the knowledge we have lately 
acquired of the bionomics of the leprosy micro-organism, they have 
only proved that experimental inoculation with small quantities of 
leprous material is incapable of producing the disease in the human 
being in every case. 

In Aiming's well-known case of the convict Keanu who was 
pardoned on condition he allowed himself to be inoculated with 
leprosy, the disease did develop, but the experiment is somewhat 
spoiled by the fact that the man had lepers among his relatives and 
was in contact with them before and after inoculation. 

(3) That in Norway leprosy diminished considerably in the 
course of sixty years, but that only about a quarter of the lepers 
were ever segregated at one time. 

This is not quite correct; it is true that only a small proportion 



58 


of all the lepers known to exist were placed under strict segregation, 
that is, in special asylums, but these were the cases unable, incapable 
or unwilling to carry out segregation at home. All the rest were 
avoiding at home all contact with their relations, etc., under 
medical supervision. In this sense the system was one of efficient 
universal segregation. 

A further similar argument, often brought forward by non- 
contagionists, is that leprosy disappeared in the Middle Ages from 
England, France, Germany, etc., without strict measures of 
segregation having been resorted to. 

It appears to me that the measures adopted to prevent the spread 
of the disease were extremely severe. Lepers were not allowed into 
churches or market places, had to carry a distinctive dress, and had 
to make their presence known by a bell or clapper. In a few words 
the intimate contact between the healthy and the diseased, which 
gives rise to the most favourable conditions for transmission, was 
most radically avoided. I also believe that plague directly and 
indirectly swept away a great proportion of the leper population, 
as they were feeble, and during times of epidemics and commotion 
would receive no alms. This observation has been confirmed to a 
certain extent in India in modem times. 

It will be seen that non-contagionist arguments are founded 
mostly on negative evidence. Positive figures bearing on the 
contagiousness of leprosy are available from Norway, Japan, and 
the Sandwich Islands, and they unanimously point to the same 
moral. 

Kitasato’s statistics from Japan show that children of lepers 
become leprous in a proportion of only 7*95 per cent, of the total. 
Matrimonial infection was proven in 3 8 per cent, cases, whilst 
persons living under the same roof contracted leprosy in a propor¬ 
tion of only 27 per cent. Brothers and sisters infect each other in 
a ratio of 4*2 per cent. These figures may need correcting 
according to the latest statistical methods. However, they roughly 
correspond to the experience gained in Norway where Sand and 
Lies’ figures differ somewhat, but show that the children of a 
leprous mother are more frequently infected than those whose father 
alone is diseased. This proves that the more intimate contact 
between mother and child leads to a greater percentage of 
acquisition of the scourge. 



59 


Sand’s statistics from Norway show that in 357 married couples 
in whicfrthe father alone was a leper, 1,241 children were born, of 
which 63, equal to a proportion of 4*9 per cent., became lepers. 
In 138 other married couples observed, the mother only was a leper; 
of 533 children born of these unions 56, or 10*5 per cent., developed 
leprosy. In 17 couples both parents were diseased; of the resulting 
63 children 8, or 127 per cent., became infected. 

Lies’ figures from South-Western Norway give somewhat 
similar results. 230 married couples in which the father only was 
a leper had 769 children, of which 79, or 10*2 per cent., contracted 
the disease. In 223 married couples the mother only was leprous; 
of 648 children, 106 of these, otherwise 16 36 per cent., became 
lepers. In 28 instances both parents were lepers; out of 79 children 
29 fell victims to leprosy, that is, 39*19 per cent. 

McCoy and Goodhues’ observations have been made in the 
Sandwich Islands, and relate mostly to cases of infection noted 
among the kokuas or voluntary helpers of the lepers at the Molokai 
settlement. Their conclusions show that : 

(1) Of 119 men, practically all Hawaiians or persons of mixed 
Hawaiian blood, living in the same house with the lepers, five, i.e., 
4*2 per cent., developed leprosy. 

(2) Of 106 women, practically all Hawaiians or persons of 
mixed Hawaiian blood, living in the same house with lepers, five, 
i.e., 4 7 per cent., developed leprosy. 

(3) Of 12 women, all Caucasians, who lived in such contact 
with lepers as is necessary in administering to their bodily and 
spiritual wants, none developed the disease. 

(4) Of 23 men living under the same conditions in contact with 
lepers, three, i.e., 13 per cent., developed the disease. 

(5) The shortest period in which the disease developed after 
the person entered the settlement was three years (two cases), and 
the longest seventeen years. 

In a report made in 1886 it is asserted that of 178 kokuas, 17 
became lepers in a year. In a later report, made in 1888, it is 
remarked that of 66 kokuas examined 23 were found to have become 
lepers. 

Such a condition of affairs does not exist at the present time at 
Molokai, and the change for the better is no doubt due to the great 



6o 


improvements made in the sanitary conditions of the settlement in 
recent years. * 

Of course it should not be forgotten that, as the kokuas are 
drawn from a population in which leprosy is not very uncommon, it 
is quite possible that a certain number of cases came to the settle¬ 
ment in a stage of incubation. 

Hollman examined carefully the conditions affecting the develop¬ 
ment of leprosy in the children of lepers at the Molokai Settlement, 
and came to the following conclusions: 

(1) It is shown that 40 per cent, of the children born of parents 
of whom one or both were lepers died under one year. 

(2) 32 per cent, of the males who were exposed ten or more 
years developed leprosy. 

(3) 4 per cent, of the females whose average time of exposure 
was less than five years developed leprosy. 

(4) 10 per cent, of the males exposed for more than seven years 
developed the disease. 

( 5 ) *3 P er cent, of the females exposed from one to seventeen 
years, and under observation seven or more years, became lepers. 

(6) The average time of exposure of the cases which developed 
leprosy was five years. 

Accordingly, the danger of contracting leprosy for children 
born of leprous parents increases with the length of exposure. 

Incidentally these investigations show that heredity does not 
play any important role in the causation of leprosy. Also, if it 
did, the disease would soon die out in any country. 

The figures and conclusions show so clearly that contagion or 
infection through immediate contact is the usual mode of 
communication, that it appears rather far-fetched to seek an insect 
carrier of the scourge. 

As a matter of fact, all experiments to prove this mode of 
transmission have so far failed, though it appears quite probable 
that the common house-fly can suck up the germs of the disease from 
open sores, carry them about for several days, and disseminate 
them in such a fashion. 

The horrid sight of flies swarming and hovering over the 
purulent sores and round the nostrils of leprous beggars is well 
known to the traveller in eastern countries. 



6i 


Graham-Smith, having shown that house-flies can harbour the 
bacteria of tuberculosis for twelve days or more, it seemed probable 
that the micro-organism of leprosy would show an equally long 
permanence in the fly-intestine. 

Leboeuf examined numerous specimens of Musca domestica 
caught on the sores of lepers, in the wards, and in houses not 
further than 150 metres from the hospital. 

He found leprotic ‘ globi' in the intestines of flies captured and 
kept for twenty-four hours, and acid-fast rods in flies thirty-six 
hours after feeding. His conclusions are that : 

(1) Musca domestica can absorb enormous numbers of Hansen’s 
‘bacilli' by nourishing itself on sores containing these germs. . 

(2) The ‘bacilli' can be found in abundance and, apparently, 
excellent condition, in the excreta of the infected house-flies. 

(3) It does not seem that multiplication takes place in the 
digestive tract of Musca domestica , but in any case there are no 
signs of degeneration. 

(4) Musca domestica possibly plays an important part in the 
dissemination of leprosy by depositing its excrements on the mucous 
membranes or small abrasions of the skin of healthy people living 
in the immediate vicinity of lepers whose sores contain ‘ bacilli.' 

It will be seen that in any case the fly does not do more than 
eventually disseminate the micro-organisms it has ingested, in a 
similar fashion to flies disseminating typhoid. 

As far as our knowledge goes, no insect plays a real role of 
transmission in any bacterial disease. Moreover, transmission 
implies a more or less complicated developmental cycle in the body 
of the intermediate host, after which a protozoon can be inoculated 
by the proboscis of a biting or stinging insect. With bacteria a 
contaminatory communication through the faeces or by the regurgi¬ 
tation of the crop contents takes place. This is the case in bubonic 
plague or typhoid. 

The fly is eminently adapted for a contaminatory or mechanical 
method of dissemination, but the difficulties inherent to the 
communication of leprosy to animals will render experimental work 
in this direction very difficult to accomplish. 

It will be seen that the intimate personal contact, as found 
between a child and its mother, gives the most favourable circum¬ 
stances for the acquisition of leprosy. 



hi 


In South Africa, where the use of coloured and native servants 
and nurses for children is very widespread, it is not astonishing to 
find that about 40 to 50 per cent, of the white (Caucasian) patients 
under segregation cannot tell where they have contracted leprosy. 
Leprosy is quite widespread in the native population. In single 
instances, after a great deal of investigation, I was able to find out 
that the disease could be traced to a coloured nurse or servant who 
had evident signs of leprosy or eventually had been segregated in 
some asylum of the Union, the disease developing in the child many 
years afterwards. 

Native labourers suffering from leprosy appear to be the source 
of contagion in many other cases. The Cape of Good Hope 
Leprosy Commission of 1895, having given careful attention to this 
matter, quotes several observations which support these views: 


‘ An individual may be leprous for many months, and it may be for a year 
or longer without showing outward signs of leprosy, and during that time may 
have communicated the disease to several without being aware of the fact that 
he is a leper himself. Hence arises the difficulty of knowing how and when the 
healthy may have been in contact with the contaminated. It is self-evident 
that the length of the period of incubation, and the possibility that during the 
period of incubation one individual may communicate the disease to another, 
greatly add to the difficulty of proving the source of contagion. Hence one may 
readily understand that statement of many lepers, that they had never been in 
close contact with a leper, or had never seen a leper, and consequently do not 
know how they got the disease. In many cases, however, possibly in by far the 
greater majority, it may be shown that at some period or other those who have 
acquired the disease have been in contact with a person or persons suffering from 
the disease ; and even where direct contact cannot be proved, indirect means 
of acquiring leprosy, such as the use of the same bedding, or articles of clothing, 
or utensils, or a pipe, or implements used by a leper, or living in a room occupied 
by a leper, or attending to a leper, may be proved to have been present at some 
time or other in the life history of those who have contracted the disease. 

In the evidence given before your Commission, Dr. A. J. J. Simons, of 
Malmesbury, who, in the course of an extensive practice reaching over a period 
of years, had many opportunities of observing the disease, gives a very instructive 
history of the course of leprosy on a certain farm in that division. In the case 
under consideration the leprosy was first noticed by him in a coloured man. a 
bastard Hottentot; next a little girl, the farmers daughter got the disease ; 
next another coloured man on the farm, a wagon driver who used to be great 
friends with the bastard Hottentot became diseased; after the wagon driver 
his master, the farmer, became leprous; next a girl who came to the farm to attend 
the farmer’s daughter and nurse her ; and finally the farmer’s niece who frequently 
visited the farm, spent the day there and was generally intimate with the leper 
daughter of the farmer. Everybody who is acquainted with the patriarchal 



63 


mode of life of the South African farmers, and who knows the habits of our rural 
population, will admit that the account given by Dr. Simons is a history typical 
of what happens in every country district in South Africa generally.’ 


* Dr. Nieuwoudt, Darling, records the following cases: (i) A patient, in 
whose family, apparently, the disease was not present, was waited on by a leprous 
servant, and developed the disease in a mixed form. Later on a boy who attended 
on him, etc., got the disease and died ; later on the gentleman’s daughter developed 
the disease in the tubercular form and is still here, though her father has since 
died. (2) An old man had the disease in whose family the disease did not exist, 
so far as known. His brother’s son, who wore the same hat of his, got the disease ; 
and another man outside this family, and of a clean family, in whose bed the 
patient slept, also developed the disease.’ 


‘ Dr. Newnham, Aliwal, knows a case where a farmer was clearly infected 
by his native coachman who had been long a leper. Master and servant when 
travelling together used the same horn drinking cup.’ 


‘ Dr. Vanes, M.L.A., of Humansdorp, writes: “ In a family living at Patentie 
there were four children affected, the parents being healthy.” With regard to 
these last cases it must be noted that many examples may be cited, which prove 
that children get the disease before the parents. Such cases appear to your 
Commission to plead powerfully in favour of the contagiousness of leprosy and 
as powerfully against its heredity. More especially in those cases in which no 
remote family taint can be shown. 

Dr. Barry, of Bedford, quotes the case of a Dutch farmer, V.H., who states 
that he got the disease through handling a spade used by a leper Hottentot.’ 

These observations might possibly be set aside by the hyper¬ 
critical if they did not agree in a striking fashion with the inferences 
deduced from the investigation of the spread of leprosy in many 
other parts of the world. 

We know that at the present moment about twenty-five to fifty 
lepers or more are living in England : and yet of these only one has 
acquired the disease in the United Kingdom (mother and father 
were lepers). The simple precautions these unfortunates are able 
to take to keep themselves separated from their families have been 
sufficient to prevent contagion. 

As a contrast we have India, where the last census appears to 
show an increase of lepers in the last ten years from 100,000 to 
110,000. 

The modern medical eye looks, therefore, upon leprosy as a 
disease which is definitely contagious, but to a very slight degree 



6 4 


under proper sanitary conditions. In situations where hygienic 
precautions are defective and the contact between the diseased and 
the healthy is unnecessarily immediate, where a leper is obliged to 
sleep in the same bed with other members of the family, and 
personal cleanliness apt to be in abeyance, the danger of contagion 
is certainly present to a markedly increased extent. 

REFERENCES 

Dehio, K. (1910). Uber die Verbreitungsweise der Lepra. Lepra XI, 16-24. 

Hollmann, H. T. (1910). Heredity versus Environment in leprosy. Public Health Bulletin 
No. 39, XI, 43-49. 

Honeij and Parker (1914)* Leprosy : flies in relation to the transmission of the disease (a 
preliminary note). Joum. of Med. Research, XXX, 127. 

Kjtasato, — (1910)- Die Lepra in Japan. Lepra, X, 144-151. 

Leboeup, A. (1912). Dissemination du bacille de Hansen par la mouche domestique. Bull. 
Soc. Path. Exotique, p. 860. Ref. Lepra, XIV, pp. 119-124. 

Lie, H. P. (1911). Statistisches uber Lepra. Archiv fur Derm, und Syphilis, CX, 473-486. 

McCoy and Goodhue (1913). The danger of association with lepers at the Molokai Settle¬ 
ment. Public Health Bulletin No. 61, XX, 7-10. 

Sand, A. (1903). Beobachtungen uber Lepra. Lepra, III, 7-16. 

- (1910). Gcschieht die Ansteckung der Lepra duch unmittelbarc Ubertragung ? 

Lepra, XI, 39-46. 


IX. EARLY AND DIFFERENTIAL DIAGNOSIS. PROGNOSIS 

The early recognition of leprosy presents difficulties commen¬ 
surate with its importance; for at times the symptoms may be so 
vague and indefinite that in the absence of conclusive microscopical 
evidence a correct diagnosis can only be made by a process of 
exclusion. Cases in which nodules containing numerous acid-fast 
rods, or in which typical maculae accompanied by anaesthesia or 
even contractures can be detected, cannot generally be considered 
as early conditions. Moreover, these ought to be diagnosable by 
someone who has a practical knowledge of the disease. It is in 
connection with slight and evanescent symptoms that serious 
difficulties arise, for such indefinite appearances are generally the 
only ones noticeable during the prodromal stage. 

Lepers with very slight and atypical lesions are not necessarily 
early cases from the chronological point of view, because the slight 
areas of anaesthesia or discoloration may have existed unnoticed 
for years. 



Sir Patrick Manson mentions the case of a gentleman who 
developed leprosy live years after noting in Ins diary a long series 
of headaches, transient fever attacks and progressive deterioration 
of health and vigour. These prodromal symptoms were followed 
three years afterwards by profuse perspiration which occurred 
without obvious cause. Such carefully observed cases are not 
frequent, and in view ot the abnormal length of the period of 
incubation of leprosy it is often impossible for the patient to 
remember the occurrence of slight indefinite prodromal symptoms. 

What in many cases is apparently the onset of the disease is 
but the more marked appearance of symptoms which have been 
present for a long time. Correspondingly very slight evidences of 
leprosy may be the traces of much more apparent previous patho¬ 
logical processes. It is accordingly often practically impossible to 
say whether we are dealing with an early case, though the symptoms 
may not be conclusive. 

It should not be taken for granted that any case showing 
leprotic symptoms necessarily will progress and become irrecog- 
nisable through deformities and mutilations because leprosy may 
get arrested in the incipient stage. 

A disease with such multiform variety of clinical features is 
naturally often extremely difficult to recognise in its first stages, 
not only because of its tendency to produce symptoms which are 
not typical for any single condition and common to several ailments, 
but also because of the corresponding variety of initial appearances. 

As a rule anaesthetic cases begin with a peculiar numbness in a 
digit, preferably the little finger of one or both hands. This 
numbness the patients compare to feeling that their fingers are 
covered with leather. On plunging the hand in warm water a 
peculiar sensation is noticed in so far as the finger or fingers affected 
feel the pain caused by the heat to a much lesser degree. The same 
symptoms may first show themselves in a little toe. On observation 
it may be seen that the fingers affected are glossier, more shiny than 
the corresponding or other fingers of the same hand. On taking 
exercise in warm weather it will be noticed that these areas are much 
less apt to perspire. Contemporary with these anaesthetic symptoms 
maculae or discoloration may be present or not in other parts of 
the body. Under some circumstances the maculae can be so slight 



66 


as to amount to little more than a peculiar discoloration very similar 
to cutis marmorata or mottled skin due to effects of cold, or they 
may take the shape of slight red lines which may form a closed 
figure. In course of time a case like this becomes a definite maculo- 
anaesthetic leper, or develops nodular features. A third alternative 
is that the symptoms may slowly but surely disappear, leaving 
hardly any trace. 

The insensibility to heat and cold may be accompanied or 
heralded by formication and shooting neuralgic pains in the limbs 
affected. Profuse sweating precedes often the anhydrosis. 

Pigmented spots may appear on various parts of the body, 
varying in colour from a light lemon yellow to a dark dirty brown. 
On looking at the spots under various angles of light incidence it 
will be noticed that the skin does not reflect the light everywhere in 
the same fashion, but is rougher in parts and appears covered with 
a slight farinaceous deposit which consists of desquamating 
superficial epithelial cells; the surface is intersected with very fine 
wrinkles, and has a dry atrophied appearance. 

Another early stage consists in puffiness and infiltration round 
the alae nasi, or thickening of the lobes of the ears and loss of the 
eyebrows, accompanied often by intensive irritation and itching of 
the neck and back of the scalp of the head. In such instances 
epistaxis and dryness of the nostrils may be followed by a 
persistent cold in the nose. 

A further early stage may be shown by a small pimple or blind 
boil, appearing in some part of the body, generally the face, this 
being followed by an outcrop of typical small nodules, superficial 
or subcutaneous. Under certain circumstances some sore or wound 
may be found to heal with extreme difficulty, or a slight abrasion 
may turn into a chronic ulcer; these have often been considered to 
form the initial sore or lesion of leprosy. They are at times, 
however, but a symptom of an infection which is already fully 
developed. 

In dealing with such extremely early cases one should never fail 
to map out the extent and measure the intensity of the paraesthetic 
symptoms, remembering that in the beginning maculae are usually 
hyperaesthetic; pin-pricks, etc., are painful which under ordinary 
circumstances would cause hardly any discomfort; the sense of 



6? 


touch can be unimpaired. Special value is to be laid on the faculty 
of recognising blunt and sharp (a penholder or a sharpened lead 
pencil will do for the purpose) and in each case a trial with two test 
tubes filled with warm (50°-6o° C.) water, and cold water should be 
used to discover the patient’s sensibility in distinguishing hot and 
cold in the affected areas. Slight muscular atrophy of the interossei 
should be looked for. 

In all doubtful cases a microscopical examination of maculae or 
discolorations should be undertaken, though the interpretation of 
any histological lesions observed is a matter requiring expert 
knowledge. Needless to say, acid-fast rods should be searched for 
in the nasal mucous membrane; they may be present for a short 
time and then disappear for longer or shorter intervals. 

Small nodules can be grouped up and treated with Anti-formin, 
centrifuged and the sediment examined for bundles of acid-fast 
rods. This method should, however, be only used in conjunction 
with an histological examination, because single acid-fast micro¬ 
organisms, without the lesions described elsewhere, are not sufficient 
to diagnose leprosy. 

The fusiform thickening of the ulnar or other nerves, which is 
one of the cardinal symptoms of anaesthetic leprosy, as a rule does 
not come into practical consideration as a help in the early 
diagnosis of leprosy, because in the initial stages it is not usually 
present to a sufficient degree to render its detection easy. Moreover, 
such swellings are often deeply seated, and therefore not accessible 
in clinical examination. 

Differential Diagnosis 

Apart from the histological and serological features which have 
already been discussed, the differentiation of leprosy from other 
skin lesions, which are somewhat similar in appearance, is generally 
based on the assumption that leprosy is an extremely slow and 
intractable disease, and that its manifestations are essentially 
indolent in character. 

Much has been written on the difficulty of distinguishing leprosy 
from syphilis. There can be no gainsaying that in very rare 
instances—a case has been illustrated by Graham Little—a 



68 


momentary confusion can arise, but a practical knowledge of both 
complaints soon shows that they must be quite different in their 
naked eye appearances. 

Syphilis and yaws as a rule produce, in all stages, lesions which 
are more vascular, angrier, with a more active and proliferating 
appearance than leprosy, which is an eminently indolent, torpid, 
slow pathological process. 

This difference is also applicable to the scars left by ulcers in 
these diseases. Syphilitic scars are usually rough, with numerous 
small fleshy warts and excrescences, and their margins are irregular 
with numerous small pockets and indentations. At times peculiar 
minute bridges of tissue can be observed under which a sound or 
needle can be passed. All these appearances are the result of a 
process of active proliferation and production of cicatricial tissue. 
Leprosy leaves scars which are smooth, shiny, and usually flush 
with or gradually merging into the surrounding healthy skin. 

Accordingly any circinate macular efflorescences in syphilis and 
yaws should be more raised, redder, bleed more easily than a 
corresponding leprotic manifestation. In addition they should 
respond to mercury or salvarsan. 

Practically every macular leper shows at least incapacity of 
distinguishing blunt from sharp and hot from cold (therman- 
aesthesia) in some area of his body. Without this crucial test it is 
not advisable to diagnose leprosy, however typical the lesions may 
appear to the naked eye, unless the opinion can be confirmed by the 
microscope. 

Leucoderma in the negro and dark-skinned races is often 
mistaken for leprosy. Here again the lack of anaesthetic or 
thermanaesthetic symptoms, the uniformity in hue and spread of 
the white spots, ought to facilitate a decision. 

Lupus and tuberculosis of the skin are the pathological processes 
whose manifestations may cause considerable difficulty. However, 
lupus is rarely multiple, has an early tendency to break down and 
ulcerate, forming scars at its periphery. Lupus is more friable and 
bleeds less easily, and though more destructive than leprosy is 
usually somewhat more amenable to local treatment. 

Pellagra ought not to be confused with leprosy, and vice-versa. 
Pellagra is symmetrical, strictly confined to parts generally exposed 



69 


to the action of the sun: there are no marked disturbances of 
sensibility in the areas affected, and the lesions are much more 
uniform and regular than is usually the case with lepra. The edges 
are not indented or irregular. In doubtful cases the microscope 
should decide. 

The ringed circinate maculae seen in Caucasians in early stages 
of trypanosomiasis are not easily distinguished from some phases 
of early macular leprosy; in fact in one case to my knowledge the 
presence of the diagnosis was only cleared up by finding trypano¬ 
somes in the blood. Leprotic lesions are, however, more permanent 
and are not as fleeting as the similar eruption in trypanosome 
disease (‘ trypanides ’). 

Syringomyelia, especially in its initial stages, is extremely 
difficult to distinguish from lepra, unless oculo-pupillar paralysis 
(consisting in myosis, diminution of the eyelid aperture and 
decreased prominence of the eyeball, Moritz) is present in the 
former eventuality, or typical maculae can be detected in the latter. 
A search for clumped acid-fast rods should be made. Early leprosy, 
moreover, does not often show such generalised lesions as 
syringomyelia. 

In advanced cases, symptoms on the part of the medulla 
oblongata, scoliosis, and the absence or presence of thickened 
nerves, ought to help the diagnosis. 

The absence or presence of thermana esthetic and analgetic 
symptoms should be carefully investigated when dealing with 
coloured races, for numerous skin diseases due to fungi cause 
lesions which resemble leprosy in quite a disconcerting fashion. 
Here, again, microscopical evidence should not be missed. 

In conclusion, it need only be added that in many instances the 
locality from which a patient comes, etc., the possibility or 
probability of contagion, are facts which, though they cannot carry 
conviction in every case, still may on the other hand simplify the 
diagnosis. It will be seen that very often recourse will have to be 
made to trained microscopical and bacteriological examination of 
excised fragments of skin. 

The extreme variability of all the initial features of leprosy 
and the chronic course of the complaint, allow only a most guarded 
prognosis to be stated in any single case of the disease. It cannot 



70 


be denied that, given skilful medical attendance and healthy 
surroundings, even this most dreaded complaint can be held in 
check for comparatively long periods. It is true that little can be 
done for advanced cases, except from the standpoint of surgery. 
Early cases, however, will show but little impairment of their 
general health under favourable conditions and arc quite capable 
of living a useful life for years and years. 

From what has been said in connection with early diagnosis, it 
is evident that it is not safe to predict in the early stages what 
course leprosy is going to take; as a rule, cases which have begun 
with indefinite anaesthetic symptoms, last longer than those whose 
nodules have appeared in batches or have shown an extensive 
appearance of simultaneous patches. 

The extensive mutilations of many anaesthetic lepers are but 
too often due to lack of attention to trifling casual wounds and 
burns. 


X. THE TREATMENT OF LEPROSY 

Not only nearly every imaginable drug, but in addition poisons 
and venoms, have been tried on lepers with the intention of 
discovering a cure, but this notwithstanding we have yet to find a 
prompt and sure method of treatment. 

The present position can, however, be summed up in the words 
that even the most useful therapeutic substance still requires the help 
of an early diagnosis and proper hygienic conditions; but that at 
no time has the outlook regarding the treatment of leprosy been so 
hopeful as at the present moment. 

In its initial stages leprosy affects the general well-being and 
appearance of an individual to such a slight extent that any thera¬ 
peutic effort which succeeds in arresting the disease in its early 
stages in a fair proportion of cases would practically amount to a 
cure. 

Spontaneous remissions of the disease, and spontaneous 
apparent ‘cures* take place, however, in a small proportion of 
lepers, and can last for several years, in some instances as long as 
15 years or more, after which the disease may again become virulent 
and rapidly carry its victim to the grave. 



7 1 


Therefore any drug or method of treatment must stand the test 
of time—five years or more—and have been carried out on a 
sufficient number of patients. 

Without taking into consideration these important factors it is 
absolutely irresponsible to speak of cure in a chronic, slow, 
intractable malady such as leprosy. 

A review of all therapeutic attempts would fill a volume by 
itself. I may, however, be allowed to give the results of the 
experiments made at Robben Island, and in South Africa generally. 

For all advanced nodular stages Chaulmugra oil, or better still 
its refined constituent 4 anti-leprol,’ injected intra-muscularly 
3-5 c.cm. at a time, is still the best palliative I know of. The 
injections should be repeated every three days and the course last 
five months or more if the patient can stand it, because at times the 
injections may become very painful. Chaulmugra oil and 
Antileprol can also be given internally in small capsules. 

Antileprol is decidedly preferable for the latter mode of 
treatment, as it does not cause the gastric disturbances which are 
the outstanding feature of the unrefined oil. Doses varying between 
15 m. and ten times that quantity can be taken daily. 

Chaulmugra oil is a product of the seeds of the Taraktogenos 
kurzii , whilst the oil extracted from Gynocardia odorata ought to 
be called false chaulmugra (D. Hooper; Article on Taraktogenos 
kurzii in ‘The Agricultural Ledger/ 1905, No. 5). It appears that 
substitution and adulteration of this oil is extensively practised, 
owing to the difficulty of keeping up a regular supply of the seeds 
of the Taraktogenos kurzii . Further particulars relative to this 
important subject are to be found in the paper by Hooper, which 
no doubt explains the variable results obtained in treating lepers 
with chaulmugra oils from different sources. 

Brocq and Pomaret mix 70 parts of chaulmugra oil with 30 of 
eucalyptus oil, and find that the injections of this product are much 
less painful than the pure oil. 

Early macular cases appear to respond better to other forms of 
treatment. A cultural extract has been prepared from Kedrowsky’s 
isolation of the micro-organism of leprosy on similar lines to Koch’s 
Alt-Tuberkulin. It contains accordingly the water-soluble toxic 
constituents of the bacillus, which, however, in the case of Hansen’s 



7 * 


germ are very feeble indeed, so that much higher concentrations 
have to be injected than is the case in the analogous treatment of 
tuberculosis by tuberculine. 

Dr. MacLeod of Charing Cross Hospital, and Dr. T. S. Davies, 
Resident Medical Officer of the Pretoria Asylum, have made use of 
this substance on single chosen cases. The results in both instances 
have been very satisfactory. The maculae or spots have slowly 
faded away. 

Dr. MacLeod's patient, a young lady from the West Indies, 
has improved to such an extent that the disease is no more 
diagnosable. I am authorised to say that other therapeutic efforts 
failed to bring about any change for the better, and that the 
patient has now been observed for over two years without any 
remission taking place. On the contrary a further subjective 
improvement has so far taken place that muscular pains which were 
present in the arm affected have disappeared. 

Dr. Sidney Davies's first patient, a little girl of years, was 
covered with dusky red patches all over her body. These faded 
away in the course of treatment extended over eighteen months, 
leaving only a slight discoloration (PI. VI, fig. 6). 

Later, six other lepers were injected with the same cultural 
extract at Pretoria, with the result that a remarkable improvement 
was brought about in two (this includes the patient already 
mentioned), marked improvement became noticeable in three, and 
slight improvement in the remaining two. 

The injections are being continued, and at the time of writing 
further progress has been made, that is to say, that the maculae 
have continued fading and in some cases have disappeared, leaving 
only a trace of faint discoloration. Details will be published in 
due course. 

It now remains to be seen whether these results will stand the 
test of time, and whether the same amelioration takes place in 
larger experimental series. 

This method of treatment does not appear to be of real 
advantage in advanced nodular cases. The injection of one or two 
cubic centimetres of the cultural extract causes a rise in temperature 
which is accompanied by the appearance of bacilli in the blood. 
This may possibly lead to dissemination of the germs of the 



73 


disease. If the individual affected is young and otherwise healthy, 
and the leprotic symptoms not marked, a course of injections 
beginning with one cubic centimetre hypodermically every three days 
may be distinctly beneficial. The treatment should be continued 
for some time even after the maculae have disappeared. 

The surgical treatment of leprosy requires most careful and 
constant attention. Fingers, toes, hands and feet are constantly 
getting septic, and prompt surgical intervention can really work 
marvels. Eye complaints, which are exceedingly frequent among 
lepers, are amenable to a certain extent to operative procedures, but 
relapses often follow even after the most skilful intervention. 

REFERENCES 

Brocq ct Pomaret (i913). Nouveau produit injectable pour le traitemcnt de la lepre. Ref. 

Lepra, XIV, 124. 

Davies, T. S. (1914). Further notes on the specific treatment of leprosy by a cultural extract. 

S. Afr. Med. Record, XII, 77-8. 

MacLeod, J. M. H. (1912). Case of maculo-anaesthedc leprosy in a woman aged 25. Proc. 

Roy. Soc. Medicine, V, 142-4. 

Serra, A. (1913). L’antileprol dans le traitement de la lepre. Lepra, XIV, 63-9. 


XI. PREVENTION, ASYLUMS AND HOME-SEGREGATION. 

SETTLEMENTS AND LEPER COLONIES 

The trend of all recent research has been to prove conclusively 
that though leprosy is communicable from the diseased to the 
healthy by direct and indirect contact, still relatively simple 
precautions are sufficient to reduce the peril of infection to a 
considerable extent. Thus the probability of the scourge rapidly 
spreading and giving rise to an extensive epidemic is certainly 
remote. On the other hand, if hygienic precautions are neglected, 
the danger for the immediate surrounding and family of a leper is 
by no means to be despised, in view of the gravity of the disease. 

The fortitude with which some nations view the slow but 
continuous increase of leprosy in their midst deserves to . be 
explained, at least to obviate its being misunderstood for some less 
admirable virtue. 

If the news spread that robbers or thugs or wild tribesmen were 
killing, torturing or maiming about five hundred or a thousand 



74 


people a year, I have no doubt that expeditionary corps, the police, 
and all the full weight of official indignation, would be employed 
to crush the criminals out of existence, and quite rightly so. On 
the other hand, an acid-fast micro-organism, against which we have 
hardly any chance of defending ourselves unless we are armed with 
the sharpest weapons of preventive medicine, is allowed to corrupt 
the bodies and souls of hundreds and thousands of new victims 
yearly without a single protest being heard. This is due to the 
fact that a biological mode of thought has not yet permeated the 
body politic, and that accordingly it is not yet sufficiently under¬ 
stood that public health means public wealth and individual health. 
We know of no other method of eradicating leprosy from a country 
than by segregation. All the critics of this method of dealing with 
the disease have no other practicable suggestion to make. It is 
true that by improving the hygienic conditions of the homes and 
lives of the poorer population, by carrying through an efficient and 
widespread sanitary inspection, leprosy could be slowly and 
definitely stamped out; but to deal with the scourge promptly 
nothing else can be suggested but isolation. The only argument that 
can be brought to bear against segregation is on the score of expense. 
In some instances parsimony and reluctance to invest money in 
public health matters have been veiled by the sentiment that it 
would be cruel to separate a leper from his family and his friends. 
Two facts are, however, overlooked. One is that when an 
individual develops leprosy in a severe and noticeable form, he 
need not trouble about his friends: they leave him of their own 
accord. The other is that consideration of the feelings of one may 
mean the misery of the whole family. In South Africa one leper 
infected in the course of time, directly and indirectly, sixteen other 
members of his family. 

It is erroneous to believe that all lepers being segregated are 
compulsorily detained, however penurious may be the conditions 
offered by an asylum. Many are but too glad to find a refuge, a 
shelter from the dreadful conditions and inhuman circumstances 
under which they had to exist in the outer world. 

The new introduction of a scheme of segregation requires careful 
preparation, because experience teaches us that the mere building 
of asylums is not sufficient to eradicate successfully the disease. 



75 


Efficient isolation presupposes adequate, skilful, sympathetic 
medical attendance in asylums, and when such is forthcoming many 
of the terrors of segregation disappear. Even in leprosy-ridden 
countries it is necessary for the medical profession to learn to 
diagnose leprosy, especially in its early stages. The public must be 
persuaded of the fact that there is no credit or advantage to be 
gained from having even only a few hundred or few thousand 
lepers hidden away in the midst of the community. 

In Johannesburg a prominent political man had a leper for 
about ten years as his gardener; the boy was repeatedly diagnosed 
as a syphilitic. The mixed case of leprosy figured in Abraham’s 
chapter on the disease in Allbutt and Rolleston’s System of 
Medicine, page 682, toured South Africa with an operatic company, 
was laid up in Johannesburg with an acute leprotic attack, was 
visited and treated by a medical man who failed to recognise the 
malady. Numerous similar instances could be quoted, all showing 
that, for a system of segregation to succeed, it needs above all the 
intelligent co-operation of the general medical public. Without it 
all efforts are of little use. 

There can be no doubt that segregation in asylums, if properly 
carried out, is in every case an expensive matter. No doubt it is 
more expensive still, in the long run, if adopted in an incomplete 
fashion, because the sum total eventually employed will have failed 
to bring about a corresponding diminution of leprosy. 

The Union of South Africa is at the present moment spending 
quite £100,000 a year on its lepers (according to the estimates for 
1913-14, £97,800), and is segregating certainly not more than half 
the number of lepers which are believed to exist in the territories of 
the Union. Under the circumstances the question arises whether 
these stupendous efforts and large sum of money would not be better 
employed in stamping out diseases which are more infective, such 
as typhoid or syphilis, and show a better chance of responding to 
treatment. 

Unfortunately we are dealing with a subject which is but rarely, 
if ever, approached without a certain preconceived view and 
sentiment. The general public, on the one hand, considers leprosy 
to be extremely contagious, shuns and has an absolutely fanatic 
terror and horror of the disease. As a contrast, the patients them- 



7 $ 


selves and their families cannot as a rule be persuaded to take the 
simplest measures to prevent an immediate spread of the scourge. 
In face of this dilemma nothing but segregation seems to meet the 
requirements of both parties. 

An alternative to isolation in asylums is home segregation, 
which, however, as a rule, fails to have the desired effect if not 
aided by constant inspection, which may become so irksome that 
many prefer to take refuge in asylums. In South Africa it has not 
been a success, though in Norway it has been possible to introduce 
the system on a large scale. 

When dealing with numerous lepers, such as are to be found 
among native populations, a system of segregation in asylums does 
not seem to be advisable on the score of efficiency, expense, and 
practicability. In these instances isolation in settlements with 
natural boundaries (a river, steep hills, on islands, etc.) appears to 
be by far the best solution. Wives can then accompany their 
husbands and the usual amemties of kraal-life (cattle, agriculture) 
need not be dispensed with. A central hospital for the infirm and 
blind, a dispensary for dressings and ambulatory medical 
attendance, a home for untainted children, should be provided, but 
the inhabitants of the settlement, whether lepers or healthy, would 
have to work for themselves; the only restriction placed on them is 
that they are not allowed to leave a certain district or province. 

Amongst natives generally, especially in South Africa, pure 
maculo-anaesthetic leprosy is by far the commoner type of the 
disease. Many of these anaesthetic cases are by no means incapable 
of work; in fact, graduated exercise and manual labour is distinctly 
beneficial to their general good health. In asylums, strapping 
young natives, with a simple contracture or a few maculae, slide 
into a life of sloth, mope about the wards and yards, and if 
re-infection can take place, certainly expose themselves to its danger 
to an unnecessarily increased extent and jeopardise accordingly 
their slender but possible chance of recovery. 

Asylums for lepers cannot be done away with; they are equally 
necessary, if not more so, than hospitals for the mentally afflicted. 
Where feasible, they should be supplemented by settlements or 
leper colonies, and in exceptional and carefully selected cases by 
home segregation. 



77 


By these means there is every reason to believe that leprosy 
would soon become extinct in any country, however a firm and 
extensive hold it may have taken, however unsuccessful previous 
half-hearted attempts of prevention may have been, however long 
the scourge may have existed in the past. 


XII. CONCLUSIONS 

Though numerous questions are yet unanswered in the pathology 
of leprosy, and many points are still under discussion, it cannot be 
denied that the researches of the last ten or fifteen years have 
opened up further fields of fruitful investigation, and helped to 
elucidate several obscure problems. 

It is true that the results of some experimental observations are 
so diametrically opposed that it appears hardly possible that both 
can be correct, at least not in their entirety. This is no new feature 
in the study of diseases whose investigation presents more than 
ordinary difficulties. Malaria is a good example. First of all 
bacteria were isolated time after tinie from patients, and it was 
sturdily maintained they were the etiological factor of the disease. 
Then the protozoa in the erythrocytes were stated to be artefacts by 
men eminent in various branches of medicine. Later on the 
plasmodia were found in marsh-water and so forth. Finally, the 
present conclusive standpoint was reached, but the path to truth 
was strewn with the thorns of controversy and deeply indented with 
the pit-falls of erroneous deductions. 

This comparison explains the present position of leprosy 
research. The exercise of sound and reasoned scientific criticism 
ought to enable us to unravel, in course of time, the delicate and 
elusive thread of positive fact from the tangled skein of mistaken 
interpretations of essentially correct, but irrelevant observations. 

The main conclusions which can be deduced from a perspective 
review of recent experimental study of leprosy appear to be 
concentrated in the following main points : 

Cultivation of Hansen's ‘ bacillus' 

Any micro-organism isolated from lepers which claims to 
correspond to the acid-fast rod seen in lesions should be identified 



78 


by injection into animals and the following production of the well- 
known histological features of lepromas. 

The culture of Kedrowsky and those similar to this type are 
the only ones to fulfil this essential postulate. 

Clinical features of leprosy. 

In addition to the nodular, maculo-anaesthetic, and mixed types, 
a further variety is recognisable, wuth thick raised patches, which 
may be more or less confluent and circinate, containing numerous 
giant cells, and very scanty acid-fast rods. 

Histological appearances of typical leprotic lesions. 

In dermal nodules and corresponding lepromas of inner organs, 
there occur numerous matted masses of acid-fast rods, intra-cellularly 
and extra-cellularly situated, very slight tissue reaction and scanty 
giant cells. 

The lesions of inner organs which show caseation, necrosis or 
numerous giant cells of the Langhans type, appear in some cases to 
be due to tuberculous complications. Guinea-pig inoculations only 
can conclusively decide the nature of such appearances. 

Maculae show the features of slight chronic irritation due to the 
action of minute doses of bacterial toxins. This, in addition to the 
external resemblance between maculae and the erythema produced 
by injecting the water-soluble contents of various acid-fast cultures 
into lepers, seems to postulate that leprotic maculae may be the 
result of a local reaction similar to the one resulting in von Pirquet’s 
and similar tests for tuberculosis. 

Rat leprosy. 

This spontaneous disease presents numerous points of resem¬ 
blance to leprosy, and may yet be found to be etiologically related 
to the human malady. 

Its experimental transmission in rodents only causes analogous 
but not identical effects to the spontaneous disease. 

Transmission to animals. 

The negative results of numerous experimental inoculations of 
human beings, and the relative low infectivity of leprosy, should 
prepare us to face very numerous unsuccessful results in dealing 



79 


with this question. It is only by injecting a large series of animals 
in various ways, and then observing them for prolonged periods, 
that deposits of acid-fast ‘bacilli’ can be produced in the inner 
organs which have the essential histological features of the leproma. 

Serology. 

Wasserman’s test is incapable of distinguishing in every case 
syphilis from leprosy. The same applies to Eitner’s test. Noguchi’s 
luetin test is negative in lepers whose disease is not complicated by 
syphilis. 

Leprotic sera often show spontaneous absorption of complement. 
The serological distinction of leprosy and tuberculosis is not 
possible in every case. Tuberculine tests may be present in lepers 
showing no clinical symptoms of tuberculosis. 

Agglutination is so far of little value owing to self-clumping of 
acid-fast bacteria and low titer of leper sera. 

Contagiousness. 

The communicability of leprosy by direct and indirect contact, 
especially under defective hygienic conditions, has been established 
by numerous repeated and independent clinical observations. 

Treatment. 

Leprosy, especially the macular variety, is subject to spontaneous 
remissions and self-cures. 

Chaulmugra oil is indicated for nodular cases; cultural extract 
for macular lepers (as far as the present experience goes). 

Prevention . 

Registration and sanitary inspection of lepers’ habitations; 
segregation in asylums, settlements, or colonies; early separation 
of children from leprous parents; betterment of hygienic conditions 
where leprosy has spread extensively. 



8o 


Fig. 

Fig. 

Fig. 

Fig. 

Fig- 5 

Fig. 6 


EXPLANATION OF PLATES I-VI 


Plate I 

Filamentary interlacing, branching, non-acid-fast germ cultivated 
from the nodule of a leper on horse-serum-nutrose-agar. Some of 
the original acid-fast rods from the leprous lesion are still to be 
seen and the acid-labile bacteria have grouped themselves in a 
peculiar fashion round the 1 globi.’ This culture was injected into 
a rat, acquired acid-fast properties, and could be then regained in 
pure culture as an acid-fast ‘ bacillus.’ Stain: Ziehl-Neelsen. 
Magnification, i.ooo diameters. 

Diphtheroid, pleomorphic, filamentary and bacillary germ, isolated 
from the nodules of a leper. It is considered to be a further develop¬ 
mental stage of the previous micro-organism. These bacteria are 
slightly acid-resisting, and regained acid-fast properties after injec¬ 
tion into animals. Third sub-culture on placental-juice-glycerine- 
agar. A slight multiplication of the original acid-fast rods may 
have taken place. Stain: Ziehl-Neelsen. Magnification, 1,300 
diameters. 

Acid-fast stage of the bacteria of leprosy which are commonly met 
with in tissues. (‘ Hansen’s bacillus,’ Mycobacterium leprae,) 
Pure culture with which all identification experiments have been 
carried out. Two months’ culture at 37°C. on fish-juice-glycerine- 
agar. From a micro-photograph. Stain : Ziehl-Neelsen. Magnifica¬ 
tion, 500 diameters. 

Section of omental gland from a spontaneously leprous rat, found 
in Ipswich by Drs. Petrie and Macalister. The similarity to the 
lesions seen in human leprosy is very striking. Stain : Ziehl- 
Neelsen. Magnification, 500 diameters. 

Nodules produced in the kidney of a rabbit by the intravenous 
injection of a culture of ‘ Moeller’s smegma bacillus.’ This lesion 
simulates leprosy in one detail only : the presence of massed acid- 
fast micro-organisms. Stain: Ziehl-Neelsen. Magnification, 500 
diameters. 

Section from the skin of a leprous rat, showing the enormous 
quantities of micro-organisms which cause the lesions. The thick, 
purple masses consist almost entirely of ‘ bacilli ’; the blue spots 
are the nuclei of the connective tissue. 

From a colour-photograph taken by Sanger-Sheppard’s method. 
Stain : Gram. Magnification, 400 diameters. 


[Plates I-IV have already been published by the author in the S. Afr . Med, 
Record (1913), XI, pp. 201-222.] 



Annals Prop. Med, & Parasitol ., Vol. IX 


PLATE I 



6 . 




82 


Plate II 

Fig. 7. Lesions produced in the liver of a rabbit after intravenous injection 
of the acid-fast culture of 4 Hansen’s bacillus.’ Similar appearances 
can be brought about by the intravenous injection of cultures of 
human and avian tuberculosis, and under circumstances by the 
inoculation of leprosy * virus.’ Stain : Ziehl-Neelsen. Magnifica¬ 
tion, 500 diameters. 

Fig. 8. Lung of a mouse which has been injected intraperitoneally with 
Kedrowsky’s strain of leprosy. The .enormous quantity of 4 bacilli ’ 
present has caused no appreciable tissue reaction of a tuberculous 
nature. Stain : Ziehl-Neelsen. Magnification, 150 diameters. 

Fig. 9. Spleen of a monkey injected intraperitoneally with Kedrowsky’s 
strain of leprosy. Also in this instance the numerous bacteria have 
not brought about a tissue reaction of a tuberculous nature. Stain : 
Ziehl-Neelsen. Magnification, 35 diameters. 

Fig. 10. Omental gland of a rabbit, which received an intravenous injection 
of Kedrowsky’s strain of leprosy. The section is teeming with 
acid-fast ‘ bacilli ’ which have not caused the death of the cells they 
have invaded, because the nuclei are still capable of staining, and 
have not appreciably altered in morphology. Stain : Ziehl-Neelsen. 
Magnification, 25 diameters. 

Fig. 11. Spleen of a mouse which had been injected intraperitoneally with 
an emulsion of Kedrowsky’s strain of leprosy. The red patches are 
groups of cells which are crammed with acid-fast rods. Similar 
lesions are sometimes seen in the spleen of lepers. Stain : Ziehl- 
Neelsen. Magnification, 25 diameters. 

Fig. 12. Portion of the mouse spleen shown in fig. 11, but under increased 
magnification. The ‘ bacilli ’ are seen to have nearly completely 
filled the protoplasm of the cells without inducing necrosis or 
caseation. No giant cells of the Langhans type are to be detected. 
These are the points which on the whole distinguish this lesion 
from one of 4 tubercular ’ nature, i.e., caused by Koch’s 4 bacillus.’ 
Stain : Ziehl-Neelsen. Magnification, 230 diameters. 



Annals Trap. Med. & Parasitol., Vol. IX 


PLATE II 




. 



9 . 


12 . ir 





8 4 


Plate III 

Fig. 1.3. Localised lesion produced in the testes of a rat by the injection of 
human leper ‘ virus.’ Stain : Ziehl-Neelsen. Magnification, 500 
diameters. 

Fig. 14. Leprous lesions in the spleen of a rat six months after injection of 
a ground-up inguinal gland from the preceding animal. Drawing 
made from three different portions of the slide. The peculiar 
hyaline increase in size of protoplasm of the cells invaded by the 
4 bacilli’ is specially noticeable. Stain : Ziehl-Neelsen. Magnifica¬ 
tion. 500 diameters. 

Fig. 15. Section through the skin of a rat suffering from spontaneous rat 
leprosy. The acid-fast 4 bacilli 9 of the disease have congregated 
round the shaft of a hair-follicle. (According to Borel may have 
been deposited there by Demodex .) Stain : Ziehl-Neelsen. Mag¬ 
nification, 35 diameters. 

Fig. 16. Small, solitary deposit of 4 bacilli ’ in the omentum of a rat, one 
month after intraperitoneal injection with ground-up 4 virus ’ from 
a spontaneously infected rat. The micro-organisms are markedly 
granular, yet juice of tha spleen of this animal produced a pure 
culture of the germ, the only one achieved out of very numerous 
attempts. Stain : Ziehl-Neelsen. Magnification, 700 diameters. 

Fig. 17. Thorax gland of a white rat injected intraperitoneally with an 
artificial culture of rat leprosy. The lesion is identical with that 
produced by the injection of 4 virus.’ Stain: Ziehl-Neelsen. 
Magnification, 500 diameters. 

Fig. 18. Small deposit of acid-fast micro-organisms detected in a nodule of 
the abdomen of a mouse injected with an emulsion of a leprosy 
culture which had been heated for an hour at 6o°C. Showing 
that, killed or alive, leprosy micro-organisms are capable of 
producing similar lesions. Stain : Ziehl-Neelsen. Magnification, 
300 diameters. 



Annals Trop. Med. & Parasitol., Vol. IX 


PLATE III 











« 



i? 


tf 

4 


* 

$ 

# 




» 4 

# 


• I 



V 






86 


Plate IV 

Fig. 19. Filamentary, interlacing, branching, partially acid-fast and acid- 
resisting germ isolated from the pleura of a man, by Birt and 
Leishman. It will be seen that it shows marked clubbed fragments 
(so-called involution forms), and also some which are definitely 
‘ bacillary’ in appearance. The resistance to the bleaching pro¬ 
perties of mineral acids varies also considerably. From a pure 
culture on glucose-agar. Stain: Ziehl-Neelsen. Magnification, 
800 diameters. 

Fig. 20. Acid-fast bacteria from the water-tap at the Lister Institute. Here 
also marked pleomorphism and variable acid-fastness is noticeable, 
but in this case the culture is not known to be pure, because the 
micro-organism was not isolated on artificial media. Similar acid- 
fast rods have been found in the scraping from a water-tap in the 
Public Health Laboratory at Cape Town and in Stockholm; 
also in mule’s dung at Robben Island. These observations show 
the ubiquity of acid-fast rods. Stain : Ziehl-Neelsen. Magnifica¬ 
tion. 600 diameters. 

Fig. 21. Section marked * Experimental Leprosy, Monkey,’ sent by Professor 
Duval, and considered by this author to be diagnosable as leprosy. 
Stain : Haematoxylin-Eosin. Magnification, 60 diameters. 

Fig. 22. True branching in tubercle-bacillus from sputum of consumptive. 

From a drawing by Mr. W. D. Severn. Stain: Ziehl-Neelsen. 
Magnification, 2,250 diameters. 

Fig. 23. Pure culture of Duval’s strain of leprosy. Its coccoid appearance 
shows it to be morphologically very similar to the ubiquitous group 
of saprophytic acid-fast micro-organisms, therefore quite different 
from ‘ Hansen’s bacillus.’ From a micro-photograph. Stain: 
Ziehl-Neelsen. Magnification, 1,100 diameters. 

Fig. 24. Mononuclear cells from the peritoneal cavity of a guinea-pig, forty- 
eight hours after injection with a chromogenic strain of an acid-fast 
micro-organism. (Duval’s 4 lepra.’) Stain : Ziehl-Neelsen. 
Magnification, 1,800 diameters. 







88 


Plate V 

Fig. i. Front and back view of an off-coloured Cape-boy, with serpiginous 
or lupoid leprosy. The presence of nerve-lesions and similar 
appearances on single other cases of leprosy, allow the inference 
that the disease is due to an infection through Hansen’s ‘ bacillus,’ 
though acid-fast micro-organisms are very scarce indeed in the 
skin-eruption. 

Fig. 2. Leprous-patch of face, apparently taking its origin from the eye, in 
a Fingo native. The borders are more raised and the surface 
rougher than is usually the case in leprosy. Similar patches are 
present on the loins of this case. No anaesthesia or paraesthesia 
detectable. 

Fig. 3. Arm of nodular leper 24 hours after intradermal injection of 10 
minims of cultural extract. There is a copper-coloured areola 
round the site of injection, in the centre of which a small sore has 
developed. 

Fig. 4. Arm of an arrested anaesthetic leper, 24 hours after intradermal 
injection of 10 minims of cultural extract. Numerous blisters; 
intense local reaction, rise of temperature to ioi° F., malaise, pains. 
All symptoms passed within three days. 




Annals Trop. Med. & ParasitolV61. IX 






yo 


F>g- 5 

Fig. 6 


Plate VI 


Diagrams of case before treatment (12.9.12). On face, infiltrated 
and thickened patches, red in colour. On legs, erythematous 
patches, reddish in colour. Over feet, purplish flush. 


Diagrams of case after treatment (6.5.14). There are faintly 
pinkish-red flat discolorations on the face. There is a flat brown 
macula on right upper arm, below the shoulder. The other yellow 
discolorations are very difficult to chart, as many of the edges are 
not well defined. There is a faintly light purple flush on the limbs, 
and the insides of the hands are a darker shade of the same. 




jhifi [) 

H A n 



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.rJU va'vj f w 



9 1 


THE METABOLISM OF WHITE RACES 
LIVING IN THE TROPICS 

I.—THE PROTEIN METABOLISM 

BY 

W. J. YOUNG 

FROM THE BIOCHEMICAL LABORATORY, THE AUSTRALIAN INSTITUTE OF TROPICAL MEDICINE, 
TOWNSVILLE, N. QUEENSLAND. 


(Received for publication 2 November, 1914) 

The effect of high temperature and a moist atmosphere upon 
the metabolism of the human body has been studied by workers in 
temperate climates, the conditions being artificially produced in a 
specially constructed chamber, and also by a few investigators who 
carried out experiments upon both acclimatised whites and natives 
living in the Tropics. In almost every case attention has been 
given to the total exchanges of the body, with the object of 
ascertaining whether the altered conditions bring about the intro¬ 
duction of any new factor into the mechanisms by which the body 
regulates its thermal equilibrium. 

The most important investigations done in the Tropics were 
those of Eijkman in 1893, who determined the calorific value of 
the everyday food of a number of white men living in Java, and 
found that the average did not differ from that usually accepted 
as a standard in Europe. Examination of the food of Malays 
also showed that the heat value per kilogram of body weight was 
not materially different from that of the white men. In these 
researches balance sheets were drawn up between the total nitrogen 
taken in and that excreted, but no investigations were made into 
the separate nitrogenous products of protein metabolism. 

A number of analyses of the nitrogenous constituents of the 
urine were made by McCay (1908) in India, but were confined to 
an examination of the urines of Bengalis, a people who conform 
to a diet with much lower protein content than Europeans. 



9 2 


In the course of a study of the effect of residence in tropical 
countries upon a working white race, it was thought that some 
information might be obtained by a fairly complete examination 
of the partition of the nitrogen and sulphur excreted in the urine. 
The significance of the different excretory substances in urine is but 
very imperfectly understood, and the effect of any departure from 
normal conditions is of interest from the possibility of throwing 
light upon the processes which constitute the nitrogenous 
metabolism of the body. 

The principal product by which proteins give up their nitrogen 
in the body is urea, and the bulk of this substance appearing in 
the urine represents the ammonia produced by deaminisation of 
nitrogenous foodstuffs. It tells us nothing as to what has 
happened to the remainder of the molecule from which it has been 
derived. 

Certain substances, however, appear to be produced by the 
metabolism of the tissues, the endogenous metabolism of the body. 
Thus, the creatinine is almost entirely derived from this source, 
since the quantity of creatinine excreted on a low protein diet is 
practically the same as when the diet is rich in nitrogen (Folin). 
Only a small fraction is due to creatine taken in with the food. 
Leathes (1907) found that during fever, when the tissue metabolism 
is known to be increased, the quantity of creatinine in the urine was 
also increased, and recent experiments on rabbits by Meyers and 
Volovic (1913) have confirmed this observation, and have shown 
that the output of creatinine is increased whether the pyrexia be 
caused by infection or by confining the animal in a hot atmosphere. 

The total quantity of sulphur contained in the urine normally 
runs parallel to nitrogen, varying with the quantity and nature of 
the protein catabolised. The bulk of the sulphur is excreted in the 
fully oxidized form as inorganic sulphates, together with a small 
quantity in combination with indol and skatol as ethereal sulphates, 
these products being the result of bacterial decomposition in the 
intestine. A small quantity of sulphur is excreted in an unoxidized 
form, termed neutral sulphur, which in all probability consists 
mainly of cystine and thiocyanates. The output of neutral sulphur 
in normal individuals was found by Folin to be independent of the 
quantity of protein in the diet, and for this reason he considered 



93 

this portion of the sulphur as derived, like the creatinine, from 
endogenous sources. 

From the experiments on creatinine mentioned above, it seemed 
possible that tissue metabolism might take place to a greater extent 
in a tropical climate, and it was of special interest to find whether 
the quantity of the substances derived from this source was very 
different from the averages in urines of residents in temperate 
climates. 

Fairly complete analyses were made of the daily urine of four 
male subjects ranging in age from 25 to 36 years, and who had 
resided in Tropical Queensland from one to four years. The 
experiments extended in most cases over a week, the total urine 
being collected and each twenty four hours’ sample analysed. The 
subjects were living an ordinary laboratory life, were allowed to 
choose their own diet, as the aim was to make the observations 
under the conditions of every day life. Moreover, most investi¬ 
gators are agreed that the knowledge of the diet being controlled 
is not without influence on the quantity of food taken, whilst if a 
fixed diet be partaken, the continued sameness affects the appetite. 
As a preliminary, therefore, the diets were uncontrolled as regards 
quantity and character, excepting in one case (protocol IV) where 
the subject was taking a milk and bread diet of known nitrogen 
content. 

In addition to these four experiments, determinations have been 
made at different times of the total nitrogen and the creatinine in 
the daily urine of five subjects, and in two cases comparisons have 
been made of the quantities of these substances excreted during the 
hot rainy season and the cooler dry season. 


METHODS 

The total nitrogen was determined by Kjeldahl, and urea by 
Folin’s (1912) potassium acetate method, uric acid by the Folin- 
Schafer method and the purin bases were estimated after precipi¬ 
tating the uric acid as recommended by Kennaway (1909). 

For the estimation of creatinine Folin’s well-known method was 
employed, but some modification was required on account of the 
high temperature of the laboratory. Although numerous papers 



94 


have been published on the conditions under which this determina¬ 
tion should be carried out, such as the quantities of reacting fluids, 
the time during which the solutions are allowed to react, the 
temperature and the extent of dilution before the readings are 
made, all such conditions have been determined for room tempera¬ 
tures about I5-20°C. In this laboratory, during the hot season, 
the air temperature during the day is rarely below 27° C. and 
generally ranges from 30 to 35 0 , whilst that of the water supply 
is usually about 30°. The conditions laid down by other workers 
were, therefore, not applicable, and had to be readjusted to meet 
the requirements. For this purpose a solution containing 
10 mgms. of creatinine in the form of the picrate (mpt. 205-206°) 
in 20 c.c. was employed, and the depth of colour produced by the 
addition of alkali and picric acid compared with a 01N solution 
of potassium bichromate in the Dubosq colorimeter in the usual 
manner. Experiments were also carried out with urine. It is only 
necessary here to give the conditions established, without going 
into the experimental details. It was found that at temperatures 
between 27° and 35° C., 10 mgms. of creatinine gave the maximum 
coloration within four minutes, whilst the colour faded if the 
reacting liquids were allowed to stand for longer than six minutes. 
The best quantities of the reagents were 15 c.c. picric acid 
(saturated solution) and 5 c.c. of 10 % sodium hydroxide, the whole 
being diluted to 500 c.c. as recommended by Folin. Within limits 
the total volume of the reacting fluids did not influence the colour, 
thus 5 c.c. of urine diluted either to 10 or 20 c.c. gave the same 
reading. The colour of the diluted mixture was found to fade 
fairly quickly, the solutions were therefore compared with the 
standard immediately after the final dilution was made. Most of 
the urines examined were very concentrated, and 5 c.c. usually 
contained enough creatinine to give a reading on the colorimeter 
between 7 and 9, the standard being set at 8. If, however, the 
reading came beyond these limits the estimation was always 
repeated with a larger or smaller quantity of the urine. The usual 
cr 1 N potassium bichromate solution was employed as a standard, 
and was carefully checked by a solution of pure creatinine picrate. 

Of the other constituents the total sulphur was estimated by 
Benedict’s method, the inorganic, ethereal and neutral sulphates by 



95 


Folin’s methods, the phosphoric acid by titration with uranium 
acetate, and the chlorine by Volhard’s method. The gravity was 
determined by Mohr’s balance. 

The full details of the analyses are given in the protocols at the 
end of the paper. 

VOLUME, SPECIFIC GRAVITY AND TOTAL NITROGEN 

Determinations of these have been made on the twenty-four 
hours’ urine at different times, each experiment lasting from four 
to ten days so as to obtain a fair average. Most of these were done 
in the hot season of the year (November to April) when the 
atmospheric humidity is high, but in two cases comparisons were 
made in the cooler season (May to October). 

The full details are given in the protocols, whilst the averages 
for each set of experiments are collected in Table I, the number in 


Table i. —Volume, Specific Gravity and Total Nitrogen of the Urine 








24 Hours Urine 

No. 

Subject 

Age 

■ 

Weight 

kilos 

Protocol 

Season 



Nitrogen 

in Grams 

VoL 

cc.s. 

Spec, 
gra v. 

Total 

Per kilo 
of body 
weight 

i 

A 

3 2 

66-8 

1 

hot 

986 

1,023 

11*69 

0*175 

2 

A 

33 

66-8 

2 

hot 

772 

1,027 

11*58 

0*173 

3 

A 

33 

66-8 

3 

hot 

1,070 

1,024 * 

14*33 

0*214 

4 

B 

2 5 

6o*6 

5 

hot 

922 

1,027 

12*41 

0*205 

5 

D 

3 ° 

63-1 

8 

hot 

7%2 

1,026 

10*83 

0*172 

6 

D 

l 

3 o 

62-8 

7 

cool 

*,437 

*,017 

12*11 

0*193 

7 

E 

36 

48-2 

11 

hot 

621 

1,023 

op 

00 

r*-> 

OO 

6 

8 

E i 

35 

48-4 

9 

cool 

>>037 

1,021 

11*69 

2*242 

9 

E 

36 

48*2 

1° 

cool 

1 , 37 2 

1,012 

10*52 

0*218 

IO 

C 

2 7 

62*2 

! 6 | 

cool 

1 

i ,3 67 

i 

1,014 

11*02 

°* I 77 


Average for both seasons, 5 persons = 

1,070 

1,021 

ii *49 

0.194 

Average for hot season, 4 persons = 

817 

1,025 

Ii*i 5 

0 

So 

00 

Eijkman's average for 19 Europeans in Java = 

1,442 

1,017 

13*04 

0*200 

Pfluger and Bohland for Europeans 

— 

— 

12*67 

0*194 

Bleibtreu and Bohland for Europeans 

— 

— 

H *93 

0*233 







9 6 


the fifth column referring to the protocol containing the full data. 
For comparison are also given the figures obtained by Eijkman for 
Europeans living in Java, as well as those for Europe found by 
Pfliiger and Bohland and Bleibtreu and Bohland. The volume of 
urine varies from day to day, but the average figures show that 
during the hot humid weather, when any exertion is accompanied 
by profuse sweating, the urines are comparatively small in volume 
and of high gravity. The volume of liquid taken in was generally 
between 3 and 4 litres a day. 

Nos. 5, 6, 7, 8 and 9 show the effect of the season on the 
excretions of the same individuals. The total nitrogen also shows 
daily variations, but the averages, 8*8 to 14*3 grams, are somewhat 
lower than that given in the physiological text-books as the average 
for Europe, 14 to 18 grams. When the nitrogen is considered in 
relation to the body weight it does not differ appreciably from that 
found in Europe. In the cases examined the daily excretion of 
nitrogen was greater in the cool season than in the hot. Part of 
this difference is no doubt due to the nitrogen lost in the sweat 
during the hot weather. Eijkman (1893) found that as much as 
076 to 1*36 grams of nitrogen per day were excreted cutaneously 
bv Malays, whilst Benedict (1906) outside the tropics, found that 
during rest the nitrogen lost in this way was about 0*071 grams per 
day, but during severe muscular labour as much as 0*13 to 0*22 
grams per hour. 

During the hot part of the year a nitrogen-balance experiment 
was carried out on subject A. The numbers, Table 2, have not 


Table 2. —Nitrogen balance on fixed diet. 


Date 

Nitrogen in 
food 

N 

[trogev Excreted 

Balance 

Urine 

Faeces 

Total 

4 - 3*14 

26*58 

17*01 

2-99 

20*00 

+ 6*58 

5 * 3*14 

26*58 

18*87 

1*50 

20*37 

4* 6*21 

6.3.14 

26*58 

19*24 

3*52 

22*76 

+ 3-82 

7 * 3*14 

26*58 

20*32 

1*82 

2214 

+ 4-44 





Total . 

+ 21*05 





97 


been included in the general averages, since the diet, which was 
restricted to milk and bread and butter, was of much higher 
nitrogen content than the usual diet. The subject increased in 
weight by 1*3 kilos during the four days, and complained of feeling 
uncomfortably full after the meals. The numbers show a consider¬ 
able nitrogen retention amounting to 21 grams, a small part of 
which may be accounted for by the cutaneous excretion. 

UREA 

The excretion of urea varies with the total nitrogen. 

The average daily outputs in the different subjects varied from 
2698 grams in A to 1479 grams in E, or 87*9% to 78*4% of the 
total nitrogen excreted respectively. The quantity of urea 
generally quoted as the average for Europeans on a mixed diet is 
30 grams per diem; the quantities found were always less than this 
amount, but when considered in relation to the total nitrogen 
appear quite normal. 


AMMONIA 

The ammonia present in the urines was quite normal in amount, 
the averages varying from 0*50 grams to 0*64 grams, or 4*8%. and 
3*6% of the total nitrogen respectively. The average output for 
all four subjects being 0 57 grams. 

CREATININE AND CREATINE 

Creatine was only present in any of the urines on one occasion, 
and then only in a very small amount (012 grams nitrogen). 

A number of determinations of the creatinine have been made 
both in the hot weather and in the cooler season. Table 3 gives 
the averages over the whole periods, given in detail in the 
protocols, each figure representing the average output per day for 
a period of several days. These averages vary between 0*47 grams 
and 071 grams of nitrogen per day, representing between 3*2 and 
6*1 % of the total nitrogen. In normal individuals in temperate 
climates the daily excretion is, according to Folin (1905), from 
3’5 to 4 5 %, whilst van Hoogenhuyze and Verploegh (1905) found 
values of 4 to 6%. Leathes observed that although during fever 
there was an increase in the actual amount of creatinine excreted, 



9 8 


there was at the same time increased nitrogen, so that the 
percentage of the nitrogen which appeared as creatinine was actually 
diminished. 

If the creatinine in the urine be considered in relation to the 
body weight, column 8, it is seen that the quantity excreted varies 
in the different individuals from 8^52 mgrms. to io'62 mgrms. 
per kilo of body weight, values which agree with the figures given 
by Folin, namely, 7 to 11 mgms. per kilo of body weight. 


Table 3. —The Excretion of Creatinine 


No. 

Subject 

Season 

Protocol 

Total N. 
grms. 

Creatinine Nitrogen 

grms. 

per cent, 
of total 

N. 

per kilo of 
body wt. 
mgrms. 

1 

A 

hot 

1 

1 i-6q 

' 

o-6i 

54 

9’ 1 3 

2 

A 

hot 

* ! 

11*58 

0-71 

6-i 

10-62 

3 

A 

hot 

3 

*433 j 

0-62 

44 

9-28 

4 # 

A 

hot 

4 

1886 

0-63 

3*2 

943 

5 

B 

hot 

5 

1241 

0-64 

5 * 

10-56 

6 

D 

hot 

8 

10-83 

o -57 

5*2 

9*°3 

7 

D 

cool 

7 

I2*II 

0*58 

4-8 

9-23 

8 

E 

hot 

11 

8-83 

o *49 

5.6 

10-16 

9 

E 

cool 

9 

j II-69 

0-47 

4 ‘* 

971 

10 

C 

cool 

6 

| I 1 -02 

0-53 

4*9 

8-52 


• No. 4 on special milk and bread diet -» 26*6 grms. N. 


As in all these experiments, with the exception of No. 4, the 
diets contained creatine, the quantities of creatinine excreted are 
probably slightly higher than that which represents tissue meta¬ 
bolism alone. 

The figures, therefore, so far as they go, supply no evidence of 
a greater creatinine output in a tropical climate, nor do they show 
any marked seasonal variation. 

URIC ACID 

The average quantity of uric acid varied from 0*42 to 0*56 
grams per day, which is quite a normal figure for a mixed diet. 



99 


PURIN BASES 

The considerable variations observed in the daily excretion of 
the purin bases are to be attributed to the tea and coffee of a mixed 
diet, since the greater part of these substances in the urine arise 
from this source. This is confirmed by the fact that with A on 
mixed diet (protocol III) these substances varied from 58 mgrms. 
to 92 mgrms. per day, whilst on a bread and milk diet (protocol IV, 
the quantity was much less and more constant, and varied during 
the next three days only from 21 to 28 mgrms. 

PHOSPHORIC ACID 

The phosphoric acid varied very considerably, the average on 
mixed diet being between 1*56 and 2*61 grams per day. The 
nitrogen to P a 0 6 ratio usually accepted for male Europeans is 
5 or 6 : 1; in every case but one the average ratio found lay between 
5*5 and 5*9, the exception showed a ratio of 71. 

CHLORIDES 

The quantity of chlorides excreted was too variable in amount 
to allow of any conclusions. 

SULPHUR 

The total sulphur when compared with the nitrogen gave ratios 
N : S of 14 to 15, this is somewhat higher than the average given 
in the text-books on Physiology, viz., 12*75, but Cathcart and 
Green (1913) during a number of determinations of this ratio, when 
different proteins were superimposed on a fixed diet, found in some 
cases numbers as high as this. 

The quantities of neutral sulphur excreted were rather high, 
representing on a full protein diet from ii*8 to 20*8 % of the total 
sulphur, and averaging from 0*250 grams to 0*455 grams per day 
as S 0 3 . The average quantity on a full diet is, according to Folin, 
about 0*25 grms. per day, or about 5 % of the total sulphur. On a 
low protein diet he found no lessening in the amount excreted, the 
quantity then representing 25 % of the total sulphur. The 
inorganic sulphates and the ethereal sulphates were quite normal in 
amount. 



100 


CONCLUSION 

The number of subjects upon which the experiments were carried 
out is, of course, too few to draw any definite conclusions, but as 
far as they go they exhibit no marked variations from the averages 
obtained in temperate climates. In every case the neutral sulphur 
was high, but whether this is due to an increased tissue metabolism, 
or not, only further experiment will show. 


REFERENCES 

Benedict (1906). Joum. Biol. Chem., I, 263. 

Cathcart Sc Green (1913). Biochem. Joum., VII, 1. 

Eijeman (1893). Virchow’* Arch., CXXXI, 170. 

- (1893). Virchow’* Arch., CXXXIII, 105. 

Folin (1905). Amer. Joum. Physiol., XIII, 66, 117. 

- (1912). Joum. Biol. Chem., XI, 507. 

Kennaway (1909). Joum. Physiol., XXXIX, 296. 

Leathes (1907). Joum. PhytioL, XXXV, 205. 

McCay (1908). Scient. Mem. Gov. India, No. 34. 

Meyers Sc Volovic (1913). Joum. Biol. Chem., XIV, 489. 

Van Hoogenhuyee Sc Verploegh (1905). Zeit. Physiol. Chem., XLVI, 415. 



PROTOCOLS 

I 

Subject A.—33 years ; weight «* 66*8 kilos ; four years in Tropics 


Date 

* 9*3 

Volume 

CC.S. 

Specific 

gravity 

Total 

nitrogen 

grms. 

Creatinine 

grms. 

As 

nitrogen 

grms. 

per cent, 
of total 
nitrogen 

Mar. 19 

'>487 

1,015 

14-20 

1*65 

o*61 

4 * 3 2 

„ zo ... 

i, 4*5 

1,013 

10*25 

**47 

o *55 

5 ‘ 3 2 

„ 21 ... 

1,030 

1,028 

* 5 * 3 1 

i* 5 * 

0*56 

3-67 

• 9 22 - 

775 

1,027 

9**7 

i *59 

0-59 

6-44 

,, 2 3 - 

690 

1,026 

9-52 

**55 

OO 

U"> 

6 

6*05 

,, H - 

*75 

1,025 

12*07 

1*56 

0*58 

4*80 

91 25 ... 

745 

1,026 

9-61 

1*89 

0*70 

7 * 3 * 

„ 26 ... 

880 

1,027 

12*12 

i *99 

o *74 

6*io 

„ 27 ... 

970 ; 

1,022 

12*93 

*•52 

o *57 

4*37 

Averages ... 

986 

*,023 

11*69 

1*64 

o*61 

5-38 


II 


Date 

* 9*3 

Volume 

CC.S. 

Specific 

gravity 

Total 

nitrogen 

grms. 

Creatinine 

grms. 

As 

nitrogen 

grms. 

per cent, 
of total 
nitrogen 

Dec 

. 16 

730 

1,029 

iz -43 

2*24 

00 

6 

6*70 

,, 

17 ... 

675 

1,028 

1**31 

*•93 

i 

072 

6*34 

,, 

18 ... 

895 

1,024 

I2*38 

2*00 

o *74 

6*oo 

,, 

19 ... 

755 

*,027 

9*60 

i-6 7 

0*62 

6*46 

,, 

20 

*75 

1,026 

12*04 

I * 9 2 

071 

5 * 9 2 

,, 

21 

820 

1,028 

ii *37 

2*02 

°*75 

6*6o 

,, 

22 

720 

1,030 

11*26 

I78 

o*66 

5*86 

,, 

23 ... 

705 

1,028 

12*24 

173 

0*64 

5* 2 5 

Averages ... 

772 

1,027-5 

i.- 5 8 

I *91 

o*71 

6 14 



























102 



Averages] 87*9 3*6 4-4 i*o | 0-5 | 2*7 76*14 7*68 17*60 







Bread and Milk Diet 




104 



Averages 81-5 4*1 5-1 1-5 0-5 7-3 76-94 8-47 14-58 




io5 


VI 


Subject C.—27 year* ; weight — 62*2 kilos ; 2} years in Tropics. 




Volume 

CCS. 

Specific 

gravity 

Total 

nitrogen 

gnns. 

Creatinine 

Date 

*9*3 

Grms. 

As nitrogen 

Percentage 
of Total N. 

July 

30 ... 

'.540 

1,011 

10 86 

1*44 

°*53 

494 

», 

31 ... 

1,040 

1,020 

13*31 

1 65 

o*6i 

461 

Aug. 

1 ... 

1,170 

1,0*4 

10*37 

1*40 

0*52 

504 

,, 

2 ... 

1,480 

*,014 

11 3 * 

**45 

0-54 

476 


3 ... 

1,100 

i,o *7 

10*00 

*‘47 

°*55 

5*48 

M 

4 ••• 

'>*55 

1,015 

10*84 

1 *46 

0 54 

5 02 

», 

5 - 

■> 43 ° 

1,014 

10*25 

1-44 

053 

5*22 


6 ... 

1,625 

1,010 

10*42 

no 

0*48 

4 * 6 5 


7 - 

1,665 

1,011 

11*87 

* *36 1 

0*50 

4.27 

Averages ... 

'>367 

*,014 

1 1*02 

*•44 

o*53 

489 


VII 

Subject D.— 30 year* ; weight — 62*8 kilos ; 1-1J yean in Tropics. 


Date, 1913 

Volume 

CCS. 

Specific 

gravity 

Total 

nitrogen 

grms. 

Creatinine 

Grms. 

As nitrogen 
grms. 

percentage 
of Total N. 

July 30 ... 

2,340 

1,011 

1372 

i*8o 

0*67 

487 

„ 31 ... 

97 ° 

1,024 

1215 

1*40 

0*52 

4-^7 

Aug. 1 ... 

1,540 

1,018 

12*28 

i*66 

0*62 

5*02 

„ 2 ... 

*, 73 o 

1,012 

12*26 

1*85 

0*69 

5*60 

3 - 

970 

1,024 

11*85 

1*48 

o *55 

465 

,, 4 ••• 

1,190 

1,019 

13*86 

i *59 

o *59 

4 * 4 * 

„ 5 ••• 

*, 3 *o 

1,015 

12*10 

*•45 

o *54 

4*45 

„ 6 ... 

*, 43 o 

1.015 

11*51 

1 *53 

o *57 

4*95 

„ 7 - 

1,950 

1,012 

97I 

1-29 1 

- 

0*48 

4*92 

„ 8 ... 

940 

1,020 

11-62 

,. 5 o 

0*56 

479 

Averages ... 

*,437 

*,o *7 

12*1 I 

**55 

0*58 

4*79 













VIII 

Mixed Diet; 63*1 kilos 



12.4.14 84*4 3*2 5-3 i*4 o-8 4*9 66-07 *3* i8 *°* 6 5 











107 

IX 


Subject E.—36 years*, 48*4 kilos; 1 to 1} years in Tropics 


Date, 

> 9*3 

Volume 

CCS. 

Specific 

gravity 

Total 

nitrogen 

grms. 

Creatinine 

Grms. 

As nitrogen 
grms. 

Percentage 
of Total N. 

July 

6 ... 

790 

1,022 

9*°4 

1*28 

0*48 

5*3 

»» 

7 ••• 

9>5 

1,023 

10-47 

1*32 

0*49 

47 


8 ... 

970 

1,022 

1179 

1*28 

00 

rf 

b 

3-9 


9 

1,185 

1.016 

10*19 

1*22 

o *45 

4‘5 


10 ... 

1,118 

1,018 

11*36 

1*22 

o *45 

4 *o 


11 ... 

i,5>o 

1,016 

11*50 

>*34 

0*50 

4*3 

- 

12 ... 

895 

1,024 

12*60 

1*28 

0*48 

3*8 


13 ... 

850 

1 026 

13*08 

1*20 

o *45 

3*4 


14 ... 

*> 37 ° 

1,015 

*4-30 

1*21 

o *45 

3 ’* 


15 ... 

770 

1,026 

12*54 

1 - 3 * 

o *49 

3*9 

Averages... 

>i°37 

1,021 

11*69 

I *27 

0*47 

41 


X 


E. Weight = 48*2 kilos. 


Date, 1914 

Volume ccs. 

Specific gravity 

Total nitrogen 
grms. 

June 25 . 

1,540 

1,011 

9*97 

» 26 . 

1,150 

*j°>3 

io*86 

» 2 7 .v 

1,460 

1,012 

io *77 

„ 28. 

'.340 

>3013 

10*50 

Averages 

*> 37 * 

1,012 

10*52 











Mixed Diet 



64-17 



























109 

THE SPECIES OF PARAGONIMUS AND 
THEIR DIFFERENTIATION* 


BY 

HENRY B. WARD 

AND 

EDWIN F. HIRSCH 
(Received for publication 23 September , 1914) 
Plates VII-XI 


CONTENTS 


Introduction . 


page 

. 109 

Historical Summary. 


. no 

Material . 


. "5 

Structure or Paragonimus ... 


. 116 

General Form . 


. 116 

Cuticular Spines. 


.. 118 

The Alimentary System ... 


. 123 

The Excretory System ... 


. 128 

The Reproductive System 


. l2 9 

General Discussion . 


. 146 

The Genus Paragonimus ... 


. 146 

Cuticular Spines. 


. 148 

References . 


. * 5 * 

Explanation or Plates 


. *54 


INTRODUCTION 

In 1895 the senior author published the first account of the 
occurrence of a mammalian lung fluke on the North American 
Continent. On various occasions since then he has contributed to 
the knowledge of these forms. In 1910 the junior author found 
some of these parasites in a host which was indisputably a native 
of Wisconsin, and became much interested in the problems 
connected with the discovery, so that when in the following year he 
engaged in graduate study at the University of Illinois this topic 
was naturally selected for his work. The present paper represents 
the results of that work and of the contemporaneous and subsequent 


Contributions from the Zoological Laboratory of the University of Illinois, No. 34. 






























no 


studies of the senior author. Unfortunately, even after very 
considerable delay, it has proved impossible to obtain all of the 
material from distant regions essential for comparison and deter¬ 
mination of all the species which are involved in the literature of 
the subject. But we feel clear that the results, even if imperfect 
still in certain details, should be laid before scientific workers in 
this field without further delay that they may be tested by those 
more favourably situated to secure material in quantities. 

HISTORICAL SUMMARY 

The earliest record of a lung fluke in mammals was published 
by Kerbert in 1878. During an autopsy held in the Zoological 
Gardens at Amsterdam, Holland, in September, 1877, parasites 
were found in the lungs of a tiger, and were sent by the Director, 
Westerman, to Kerbert, who determined them to be undescribed 
trematodes and named the species Distoma westermanii. Three 
years later Kerbert received specimens, through Bolau, from the 
lungs of a tiger that had died in the Zoological Gardens at 
Hamburg, Germany. Upon this material, which proved to be 
identical with the former, he completed an extensive morphological 
study (Kerbert, 1881) of the species. 

The first information concerning the occurrence of such a 
parasite in man came from Manson, who in a letter to Cobbold, 
published with comments by the latter, described (Manson, 1880) 
the discovery of distome eggs in the sputum of a Chinaman suffering 
from hemoptysis. At the same time he mentioned a fluke found in 
another patient, a Portuguese, by Dr. Ringer. The specimen was 
sent to Cobbold, who in the note just cited named it Distoma 
ringeri. Although the description is scanty, the records leave no 
doubt as to the identity of the species. It should be noted that 
this is the first published record of this human parasite, ante¬ 
dating by nearly two months that of Baelz, to which priority is often 
accorded. 

The latter record came thus almost simultaneously, but from 
Japan. As early as 1878 Baelz had noted there conditions, similar 
to tuberculosis and regularly attributed to it, which he regarded as 
resulting from some parasitic infection. He sent the sputum to 



Ill 


Leuckart for examination and described the bodies in it as psoro- 
sperms, but before receiving Leuckart’s diagnosis, came to the 
conclusion that they were in reality eggs of some fluke. This 
opinion was first published by Manson (1882), who stated that after 
having examined some of Baelz* gregarine material he had found 
bodies identical with the ova of Distoma ringeri. In the following 
year Baelz (1883), confirmed in his view of the nature of the material 
by the report of Leuckart, published a description of the worm, which 
he re-named Distoma pulmonale . According to Stiles and Hassall 
(1900, p. 567), certain Japanese investigators in the interval had 
named the species Distoma pulmonis . Both this name and that of 
Baelz are ante-dated by the name of Cobbold, which is apparently 
the first designation used for the human parasite. There is no 
need to review here the important clinical and pathological data 
submitted by Manson, Baelz, and later writers. The wide and 
abundant distribution of the parasite in Japan, Formosa and Korea, 
its occurrence among all ages and classes of society, and the high 
ratio of the infected in certain districts are well-known and firmly- 
established facts. The Asiatic lung distome is one of the most 
important human parasites. 

The next important step in the history of this form was taken 
by Leuckart, who in conjunction with his student Nakahama made 
a careful study of specimens of the fluke which Baelz had sent him 
and of specimens of Distoma westermanii from Kerbert. Although 
he noted points of disagreement, these were interpreted as 
unimportant minor differences, and the specific identity of the two 
forms was definitely asserted. We do not find that this identity 
has been questioned since then, except once (Ward, 1908). Stiles 
and Hassall (1900, p. 561) voice the general opinion in stating that 
the human parasite ‘ though originally supposed to represent a new 
species is now generally admitted to be identical with Kerbert's 
form from the tiger.* 

Regarding the genus, however, several investigators commented 
independently on the impossibility of retaining this form in the old 
group Distoma ; and finally in 1899 Braun made it the representa¬ 
tive of a new genus to which he gave the name Paragonimus The 
first careful and extended description of the genus came from 
Looss (1899), who unaware of Braun's publication gave the form 



112 


another name that, because of its slightly later appearance, must 
be relegated to synonymy. However, the description given by 
Looss (1899) still ranks as the most accurate and complete avail¬ 
able. Even at that time Looss recognized only a single species, 
to which he gave the name Paragonimus westermanii * 

Except for its sporadic extra-limital occurrence in hosts that 
were known to have come from the regions where the parasites are 
endemic, the species was not recorded from the western continent 
until one of us (Ward, 1894) published a description of a similar 
form obtained from a cat in Michigan. This paper called attention 
to the differences between these worms and the description given by 
Kerbert and Leuckart for Distoma westermanii , but assigned them 
tentatively to the same species. It also noted the possibility that 
the host had been brought as a pet from the East, and that hence 
the parasite was not en'demic. Grave doubt was thrown on this 
view through the discovery by Kellicott of worms in the lung of a 
dog from Ohio, which were sent to Ward and pronounced by him 
identical with those he reported from the cat. 

All possibility that the parasite had been found only in hosts 
introduced from abroad disappeared when Stiles and Hassall 
(1900) recorded the abundant discovery of a similar parasite from 
hogs slaughtered at Cincinnati, Ohio. Their paper gives not only 
a very careful account of these specimens but a detailed comparison 
with other records and the best critical summary of the literature 
on these lung flukes which has been published to date. While 
recognizing the possibility that their worms ‘ may represent a 
distinct variety at least,’ Stiles and Hassall 1 feel compelled to 
continue for the present to look upon the American form as identical 
with the Asiatic.' 

Since then the lung fluke has been reported in numerous cases 
from North America. In the human host it was first diagnosed by 
Mackenzie, in 1904, from the lung of a Japanese fisherman on the 
Columbia river. Then Fehleisen and Cooper (1910) reported a 
case in a Japanese worker in California fruit orchards who had 
come to the country some six years before. Microscopic examina¬ 
tion of the sputum showed in every field two or three eggs of the 
distome. No details of size or structure are given, but the case was 

• Looss wrote P . westermanni which is evidently a typographical error. 




n 3 


of long standing and had been diagnosed in Japan as lung fluke. 
Thus all the evidence goes to show that it was not contracted in the 
United States. In the same year Null (1910) recorded its 
occurrence in a Korean at Seattle, Washington. 

From other hosts there are several cases on record. Null stated 
casually that it occurs in dogs and cats from the Oriental quarters 
of San Francisco, but gave no further data regarding the parasites. 
Nickerson (1911) recorded the occurrence of P . kellicotti in the lung 
of a cat from the grounds of the University of Minnesota at 
Minneapolis. Only three specimens were found. 

The junior author of this paper (Hirsch) observed several cases 
of marked significance in Wisconsin. As early as 1907 he noticed 
a peculiar cough in a pet cat, which became more seriously affected 
and was chloroformed a year or more later. These worms were 
found in the lungs. A kitten born to the first cat was affected in 
the same way and found to be infected with the lung fluke when 
only a little more than a year old. In addition to these cases, of 
which the full case history, written by Hirsch, was published by 
Hanson (1911, p. 112), the former has also found the lung fluke in 
two other cases hitherto unpublished. These cases are another 
kitten of the same litter and a fourth cat, unrelated and living some 
miles distant from the first three. It is positively known that the 
first three were born in that locality and had never been away from 
it, hence the endemicity of the parasite can only be questioned on 
the ground that possibly the means of infection—a fish carrying the 
encysted young distome for instance—was brought in from a 
distance. There is no probability that the two young cats were 
infected directly from the mother cat, but in all likelihood through 
the consumption of the same food. It seems improbable that all 
three could have been infected at the same date, and certainly the 
fourth cat did not acquire its parasites from the same food. These 
cases are the first in which the presence of the lung fluke was 
diagnosed in the living animal and the diagnosis confirmed by the 
demonstration of the ova in the sputum. They are also the first 
on this continent in which the place of birth and infection was 
positively determined for the host. 

The range of the human lung fluke was further extended by the 
work of Musgrave (1907). He studied seventeen cases with eight 



autopsies in the Philippine Islands, and gave a careful morpho¬ 
logical description of the parasite. His data will be utilised later 
in our paper. 

One of the seventeen hosts was a Chinaman, two were Japanese, 
and the other fourteen were native Filipinos, so that the species is 
undoubtedly endemic in the Islands. Various later papers contain 
casual references to the occurrence of this species in the Islands, 
but do not give data of value for our purposes. Garrison and 
Leynes (1909) studied the development of the ova of Paragotiimus 
in the Philippines. They do not give a precise description of the 
ova, but apparently these were obtained from cases among those 
recorded by Musgrave (1907) which have already been considered. 
The work of Garrison and Leynes, which deals with the experi¬ 
mental development of Paragotiimus ova under varying conditions 
of temperature, light, salinity and desiccation, has bearings of great 
importance on the dissemination of the disease and the infection 
of man, but does not throw any light on the problem of species and 
their differentiation. 

In a paper published several years ago, the senior author (Ward, 
I 9°3) commented on the great discrepancies between the measure¬ 
ments recorded by various investigators for the eggs of Paragotiimus 
westermanii , and gave an outline sketch representing these differ¬ 
ences in a graphic manner. He reached the conclusion that all 
records could not be accepted as correct unless more than one species 
was concerned. The measurements given by Yamagiva, which were 
made from ova in sections of the brain and lungs of man, have 
since been shown to belong, in all probability, as Ward stated later, 
to the Japanese blood fluke, Schistosoma japonicum , and are 
consequently eliminated from the present discussion. In a later 
paper (Ward, 1908) these records were discussed at greater length 
in the light of further evidence, and the conclusion was reached that 
the American form originally identified as Paragotiimus westermanii 
is undoubtedly a distinct though closely-related species, and to it 
Ward gave the name of Paragonimus kellicotti. He also indicated 
the probable specific independence of the Japanese form on which 
further work was then being done. This form, which will be 
fully discussed in the descriptive section of this paper, must bear 
the name of Paragonimus ringeri (Cobbold, 1880). 



MATERIAL 

The junior author made a detailed and careful study of 
Paragonimus kellicotti on new material obtained from the pig at 
the Cincinnati (Ohio) abattoirs. The specimens were taken alive 
from the lungs, and after shaking in normal salt solution according 
to the method of Looss, were preserved with great care. The 
technique involved nothing unusual, and gave good results 
throughout. Total preparations and serial sections of the 
specimens originally obtained by Ward from the cat in Michigan, 
by Kellicott from the dog in Ohio, and by Hirsch from the cat in 
Wisconsin, were compared item by item with this new American 
material. 

Several specimens of the Japanese form from man were also 
available for comparative study through the courtesy of Dr. S. 
Uchida, of Tokyo, and eggs from the Seattle case were kindly 
sent us by Dr. Null. This form has been described in detail several 
times, notably by Katsurada (1900) and Kubo (1912). Despite 
this fact, the structural features are not even yet well known, and 
in most respects these descriptions are couched in such general 
language that we cannot determine from the paper more than 
generic features regarding certain organs. The last paper especially 
(Kubo, 1912) falls short of what might be wished. In spite of a 
rich supply of fresh material he gives very little more information 
on the structure than the work of Katsurada a dozen years earlier. 
The figures are distinctly unsatisfactory, being in some cases vague 
and almost illegible, and in others highly diagrammatic. They 
are certainly inferior to those given by Katsurada. 

Our work was supplemented by comparison with three co-types 
of Distoma westermanii kindly placed at our disposal by Professor 
Kerbert and now in the Ward collection. While this supply was 
adequate for the determination of the most important features in 
the anatomy, as will appear in the following pages, it did not 
suffice for a complete study of the structure, and a detailed 
comparison of this form with those from man in Japan, and from 
cat, dog, and hog in North America, must be left to some future 
student who has at his command a larger supply of material. 

Despite persistent effort, it has been impossible to secure for 
study and comparison any material from the Philippine Islands, 



ii6 


so that the status of that form could not be tested by the method 
worked out on the other material. Although only a single paper 
has been written on the anatomy of this form, that one is so 
carefully worked out that it furnished very definite material for 
comparison with the work on other species. In spite of this, there 
are several points on which we would fain have had precise infor¬ 
mation concerning the structure of some organ that was not treated 
in extenso in the text of that paper. 


STRUCTURE OF PARAQONiMUS 

General Form 

Paragonimus kellicotti has a somewhat elongated form, 
elliptical in dorsal aspect. The dorsum is strongly arched, the 
highest point being somewhat anterior to the middle portion of 
the body, while the ventral surface is slightly flattened. The 
anterior end rounds off gradually, but the posterior extremity is 
attenuated, and sharply curved. The range in size is given in 
comparison with Paragonimus ringeri and Paragonimus westermanii 
in the following table: — 


Species 

Host 

Length in mm. 

Width in mm. 

Thickness in 

mm. 

Authority 

Max. 

Min. 

Av. 

Max. 

Min. 

Av. 

Max. 

Min. 

Av. 

P. kellicotti 

Hog 

I 1*0 

8-5 

9-8 

3*5 

3 *o 

3 ‘i 

3*2 

2*2 

2*6 

Hirsch 

P. kellicotti 

Cat 

1 5*5 

11-2 

136 

77 

4-8 

— 

— 

— 

— 

Ward 

P. kellicotti 

Hog 

i4-o 

3 *o 

— 

4 -o 

2*0 

— 

— 

— 

— 

Stiles 

P. kellicotti 

Dog 

20*0 

i 5 *o 


— 


— 

— 

— 

— 

Kellicott 

P. zvestermanii 

Tiger 

9-0 

7 *o 

— 

6*o 

4 -o 

— 

4 -o 

2*0 

— 

Kerbert 

P. ringeri ... 

Man 

10*0 

8-o 

— 

6-o 

V° 

— 

— 

— 

— 

Leuckart 

P. ringeri ... 

Man 

lyo 

7 ** 

9-6 

7*5 

5 '° 

5 *° 

4 -o 

3*5 

37 

Katsurada 

P. ringeri ... 

Man 

'7 

75 

io-i 

6 

4*5 

5*8 

5-5 

3*5 

5 

Kubo 







11 7 


From these figures it may be noted that there is considerable 
variation in the size of the different individuals. Taken as a 
composite picture, however, a description of the American form 
emphasizes its much elongated, relatively slender structure, which 
is in contrast with Paragonimus ringeri and Paragonimus 
westermanii> both of which are oval-shaped, thicker, and broader 
parasites. 

The most recent and extensive study of fresh material for 
Paragonimus ringeri has been made by Kubo (1912). He has listed 
measurements of thirty-six specimens from the lungs of the dog in 
Japan and of eleven specimens from the human lung in the same 
region. There seems to be no contrasted difference between 
specimens from the two hosts. The entire series averages in 
length 10*08 mm.,* in width 5*8 mm., and in thickness 5 mm. The 
smallest he found measured 3 mm. in length and the largest 12 mm. 
Worms with a length of 5 mm. and over are sexually mature, 
having some eggs in the uterus. He has not included these 
extremely small and presumably young specimens in his averages, 
so that the latter represent fairly well full-grown adults. Yet even 
with that, the omission of five conspicuously small specimens listed 
in his table will raise the size averages appreciably. 

,On the other hand, when one examines a group of these parasites 
with the eye it is not difficult to see a general difference between 
the species in type of form. Yet at the same time it is clear that 
neither the size nor the form gives a safe basis for distinguishing 
the species. There is no doubt that some of the conflicting views 
in regard to size are due to the measurement of specimens at 
different ages and stages of growth. 

At the anterior extremity is found the oral sucker directed 
towards the ventral surface at an angle of about 45 0 , while the 
acetabulum lies on the same surface in the median line slightly 
anterior to the centre of the body. In ten specimens these suckers 
average respectively 075 mm. and 0*83 mm. in diameter. The two 
suckers of Paragonimus westermanii are of about equal size, 
078 mm. in diameter, according to Leuckart, while those of 
Paragonimus ringeri are unequal, and smaller, the oral sucker 


Not io-8 mm. as given by Kubo in the text. 



being 0*53 mm. in diameter and the acetabulum 0*6 mm., or at 
most 075 mm. according to Leuckart. 

Kubo says that the oral sucker of Paragonimus ringeri in a 
medium-sized worm (10 mm. long) measures 075 mm. in diameter, 
and that the ventral sucker in a similar specimen reaches 0 8 mm., 
being thus slightly larger than the oral sucker. These measure¬ 
ments differ from those of Leuckart and agree closely with those 
we report for Paragonimus kellicotti , but here again the character 
varies so much with the size and age of the specimen measured that 
no dependence can be placed upon it in determining the species. 

The genital pore in Paragonimus kellicotti lies just behind the 
acetabulum, medial or a little to the right or to the left. The 
position of the genital pore as given by Stiles for other species is 
the same. The cuticula covering the entire body is relatively 
thick, and armed with spines (PI. X, fig. 18). The thickness of 
this structure, however, is not uniform over the entire body. 
Around the suckers it may be 0005 mm. thick, while over the 
posterior extremity it may be 0*048 mm. There is often also a 
distinct difference in the appearance of this layer. Sometimes it 
seems perfectly homogeneous, but in other places it appears as if 
made up of two distinct layers, an outer more dense and less 
refractive, and an inner vertically striated, rather clear, or highly 
refractive with numerous coarse granules. The cuticula shows no 
cellular structure, but is sharply marked from the underlying 
muscles by a distinct basement membrane. 


Cuticular Spines 

Since a preliminary study of the spines revealed characteristics 
that appeared to be of great value for the differentiation of species, 
these structures were subjected to a most precise examination, with 
results that we believe justify the labour. It should be recalled 
that Kerbert, Leuckart, and Stiles differ in their statements 
concerning the distribution of these structures and the place where 
the largest are to be found, but so far as we can ascertain no one 
has noted differences in the form of individual spines or in their 
grouping. An examination of the three worms with reference to 



these specific features gave pictures interesting both for their 
resemblances and for their striking differences. The conditions in 
the parasite from the lung of the pig are given first. 

The spines of Paragonimus kellicotti lie in irregular circular 
rows over the body and are set firmly in the cuticula with their 
free ends directed posteriad. They extend entirely through the 
cuticula, and sometimes into the body musculature beneath (PI. X, 
fig. 18). Structurally, the spines are thin chisel-shaped scales, 
usually several times as long as wide. The surface directed away from 
the body is slightly convex, while that directed towards the body 
is correspondingly concave. The free end is rather deeply serrated 
into a number of very sharp teeth. These certainly aid the 
parasite materially in effecting its movements, and in maintaining 
its position in the tissues of the host. The basal end of the spines 
is broader and thicker (0*005 to 0*010 mm.) than the tip, often 
appearing cleft so as to show in cross-section groups of closely 
aggregated oval or irregularly shaped chitinous bodies. 

There is a distinct difference in the distribution and size of the 
spines (PI. VIII, figs. 5-11) in various parts of the body. The suckers 
are entirely devoid of these structures, while the cuticula closely 
surrounding shows the transition stages between the non-spined and 
the spined condition. This holds true especially for the region 
surrounding the acetabulum. Here the body for a small distance 
around the sucker may be entirely free from spines. When they 
appear, they are few in number but very sharp and even decidedly 
hooked (fig. 10). Around the oral sucker the spines (fig. 9) are 
short with broad bases and sharp tips, a condition which is soon 
replaced by larger spines with broad serrated tips. The spines are 
set relatively more thickly over the anterior half of the body 
(fig. 5) than over the posterior (fig. 7). The cuticula on the dorsal 
surface just behind the oral sucker, and on the ventral surface 
between the suckers (fig. 8) and just behind the acetabulum, is 
especially well armed. The spines over the dorsal surface show 
little variation except in number and in length. On the ventral 
surface the greatest variation occurs around the suckers. This 
divergence, however, is not one from the characteristic type, but 
rather is a gradual diminution in size. The spines on this surface 
of the body are perhaps shorter and broader than over the dorsum. 



120 


The results of an extensive series of measurements of the spines 
in Paragonimus kellicotti have been brought together in the form 
of a table or summary. This shows the exact size of individual 
spines in various regions of the body, their distance apart, and 
their variation in those cases where marked differences occur 
(Table A). 

Table A.—Spine Measurements of Paragonimus kellicotti 



Distance apart 1 

Length 

Base 

Tip] 

Ventral Surface— 

Near Oral Sucker. 

0*005— Q, oo8 mm. 

o*oio mm. 

0*005 mm. 

o*ooo mm. 

Between Suckers. 

0*013—0*026 mm. 

0*046 mm. 

0*013 mm * 

0*013 mm. 

Acetabular Region. 

0*030 mm.—variable 

J 

0*010—0*020 

o*ooi mm. 

o*ooo mm. 

Between Acetabulum and Anterior End 

0*026 mm. 

mm. 

0*031 mm. 

0*013 nim * 

0*008 mm. 

Between Acetabulum and Posterior End 

0*013—mm. 

0*030 mm * 

0*023 mm. 

0*021 mm.} 

Posterior Extremity . 

0*026—0*047 mm. 

1 

0*036 mm. 

0*015 mm * 

0*008 mm. 

Dorsal Surface — 

Anterior Extremity . 

0*008—o*oi2 mm. 

0*031 mm. 

1 

o-oio mm. 

0*008 mm. 

Anterior One-Fourth . 

0*012—0*026 mm. 

0*034 mm. 

o*oio mm. 

0*008 mm. 

Middle of Body . 

0*015— O‘oz6 mm. 

0*044 mm * 

o-oio mm. 

0*008 mm. 

Posterior One-Fourth . 

0*026—0*036 mm. 

0*040 mm. 

o*oio mm. 

- 0*008 mm. 

Posterior Extremity . 

o*oi2 -0*026 mm. * 

0*036 mm. 

o-oio mm. 

0*008 mm.’ 


The spines of Paragonimus kellicotti are of one general type, 
chisel-shaped, with distinctly serrated or saw-toothed free 
extremities. There is, to be sure, a variation in their length, but 
in general the characteristic features of this type of spine are 
retained throughout. The study of these spines involved their 
examination over the entire body of the parasite; in no place was 
there any marked divergence in form, and nowhere did they occur 
other than singly. While the substance of the comparison was 
made upon material taken from the hog, further data were obtained 
by examining a piece of the cuticula from a parasite removed from 
the lungs of a cat. The shape of these spines, as well as their 





121 


arrangement, was the same (PI. VIII, fig. n), the only difference 
noted was that in extreme cases they were somewhat longer 
(C078 mm.), and their free ends more deeply serrated. Such 
differences may easily be due to the age of the individual parasite. 
Our material was not extensive enough to enable us to test the 
question of a possible growth in the size of these spines during the 
life of the adult parasite. But we think all will grant that this 
extreme of variation in the cuticular spines is not great enough to 
justify regarding them as separate and distinct from the type 
found in the hog, much less to conceal their essential agreement 
with that type. Their radical dissimilarity with the other types 
will appear conspicuously when those types have been described. 

The dimensions of the spines of Paragortimus westermanii, as 
given by Kerbert, have elicited considerable comment, for he 
reports that the largest spines were o‘oi8 mm. long by o - oo2 mm. 
broad at the base, and the others were o'oio mm. long. He 
describes the spines as lancet-shaped. Kerbert’s figure shows the 
posterior end of the parasite and the surface for a short distance 
around the oral sucker entirely devoid of spines. Upon 
re-examining the co-type specimens of Paragonimus westermanii 
in the Ward Collection, these statements were found to need some 
emendation. The spines are indeed lancet-shaped, but are 
considerably longer than Kerbert’s dimensions indicate. They are 
fully as long as those in the cuticula of Paragonimus kellicotti, 
being 0‘047 to 0^049 mm. long by 0 005 to 0 010 mm. broad at the 
base. In addition, spines cover the entire body excepting the 
suckers, as in Paragonimus kellicotti. Leuckart gives the measure¬ 
ments of the longest spines of Paragonimus ringeri as 0 06 mm. 
with a base of 0 014 mm. broad. These dimensions approximate 
the size condition revealed in Paragonimus kellicotti by our 
investigations. 

To judge from these data one would be forced to maintain that 
there was no clear difference between the spines in the three species, 
since all of the measurements fall well within the extremes we 
found in Paragonimus kellicotti. And yet when one takes into 
account the form of the individual spine and the arrangement of 
the spines on the surface of the body there are evident and 
distinctive differences of a striking character. The measurements 



122 


already given show that the spines of Paragonimus westermanii 
and those of Paragonimus ringeri do not differ in size much, if at 
all, from those of Paragonimus kellicotti, In the first two species, 
however, the spines are markedly lancet-shaped, while in the last- 
named they are, as already stated, broad and chisel-shaped. The 
condition in Paragonimus ringeri is different. Over certain parts 
of the body, especially on the ventral surface behind the 
acetabulum, the spines are short with broad bases and tips. Here 
they are set very closely together, forming at times almost 
continuous circular rows. Along the side of the body the spines 
are slender with sharp free ends. 

While these differences in shape are very striking characteristics, 
the spine arrangement is even more distinctive. The spines in 
Paragonimus westermanii lie rather sparsely scattered in more or less 
incomplete circular rows (PL VIII, fig. 13). Their free ends extend 
very far beyond the cuticula, giving the parasite a thorny appearance. 
In Paragonimus ringeri (PI. VIII, fig. 12) the general appearance 
is very different, since the spines are characteristically arranged in 
groups which are often massed together, and in certain parts of the 
body are close enough to form almost continuous circular rows. 
While the shape of the individual spines in Paragonimus ringeri is 
subject to some variation, in fact offering a transition series 
between Paragonimus westermanii and Paragonimus kellicotti , the 
group arrangement affords a striking and constant means of 
differentiation between the species. 

In his description of the Philippine form, Musgrave (1907, 
p. 32) discusses the distribution of the spines at length. He says 
that on the ventral surface between the acetabulum and the oral 
sucker they are ‘ almost entirely absent,* and that when they occur 
in this region they are 'smaller than the others,* whereas they 
increase in size on the lateral borders and reach a maximum on the 
dorsum. This does not agree with conditions in the forms we have 
studied and have described in the foregoing, as is shown distinctly 
by a comparison of this statement with our figures. Musgrave 
speaks of these spines as ‘scale-like,* a term difficult to compare 
with our findings, and he gives no illustration of these structures. 
From the evidence at hand we are unable to reach a final 
conclusion, but it favours the separation rather than the amalgama- 



123 


tion of these forms with any known species. The decision of this 
matter must await the study of material from the Philippine 
Islands, which as yet we have been unable to secure. 

In regard to the Japanese form, Kubo gives no data that can 
be compared with our findings, but in one part of this description 
he says that in larger and older worms the spines lose their sharp 
points and become stumpy. This might suggest the interpretation 
of the conditions we have described as possibly changes with 
growth and use, dependent thus merely upon the age of the 
specimens. This view is hardly tenable, since our illustrations are 
taken from specimens of the same approximate size, and presumably 
of the same age, if all were a single species. Furthermore, this 
explanation does not touch the striking differences in the arrange¬ 
ment of the spines on the surface of the body. 

The differences in form and arrangement of the cuticular spines 
in the species we have studied may be summed up in tabular form 
as follows: 



P. westermanii 

P. ringeri 

P. kellicotti 

Shape ... 

Lancet-shaped 

Chisel-shaped 

Chisel-shaped 


Very slender 

Moderately heavy 

Heavy 

Distribution. 

Sparsely, somewhat 
irregularly, singly 

Circular rows, in 
groups 

Circular rows, singly 


The Alimentary System 

The opening leading from the oral sucker into the pharynx in 
Paragonimus kellicotti measures 0*050 to 0*075 mm - diameter. 
It becomes somewhat larger toward the outer margin of the sucker, 
and in sectional view appears wedge-shaped. It is continuous 
proximally with a thin lamella that forms a small pocket, the pre¬ 
pharynx, which is located between the pharynx and the sucker and 
serves to unite the two. No such region is mentioned by other 
authors, and yet may be found in Paragonimus westermanii or 
Paragonimus ringeri on careful examination. 

The entrance into the pharynx is provided with four lip-shaped 








124 


projecting folds which effectively close the canal during contraction. 
The two which lie laterally are larger than those bounding the tube 
dorsally and ventrally, thus making the entrance into the pharynx 
appear much like a vertical slit. The pharynx is spherical in 
shape, about 0*4375 mm. in diameter. Its walls are composed of 
heavy muscles, and the canal itself is limited to a narrow vertical 
slit. The inner lining of the pharynx consists of a thin layer of 
the cuticula, o*oi to 0*02 mm. thick. Towards its union with the 
oesophagus the opening through the pharynx widens considerably. 

The oesophagus is nearly circular in trans-section, and about 
0*21 to 0 255 mm. long. The anterior part of the alimentary canal 
is directed towards the dorsal wall, but not very acutely. After 
the junction of the pharynx with the oesophagus this direction is 
continued, and becomes a little more abrupt. At first the 
oesophagus is about 0 08 mm. in diameter, but towards its 

branching this dimension decreases slightly and then increases 
again to 0*16 or even 0*175 mm. The thickness of the wall varies 

in this short distance from about 0*012 mm. near the pharynx to 

005 mm. somewhat nearer the bifurcation. This difference 
necessarily causes the inner lumen to vary inversely. Such 
relations as have been described are subject to variations no doubt 
due to the contraction of these parts. Structurally the oesophagus 
(PI. XI, fig. 23) is relatively simple, consisting of an inner homo¬ 
geneous layer, often thrown up in folds, and an outer well- 

developed muscular layer. The latter is made up essentially of the 
inner circular fibres, while the outer longitudinal fibres surround 
the wall in relatively heavy parallel bands. This portion of the 
alimentary canal is enveloped by well-developed glandular cells 
whose ducts open into the oesophagus, in all probability furnishing 
a salivary or digestive secretion. 

„ The two lateral branches of the oesophagus are 0*175 to 
0*255 mm * l° n g> an d are sharply differentiated from the intestinal 
caeca by several distinct characters. The intestinal caeca 
immediately attain a diameter of 0*315 mm., whereas the 
oesophageal branches measure only 0*05 mm. in this dimension. 
More generally speaking, there is in this region a sudden increase 
in the diameter of the caeca to about four or five times that of the 
oesophageal branches. In addition, the intestinal epithelium is 



125 


characterized by tall columnar cells (PI. X, fig. 17) which contrast 
strikingly with the lining of the branches of the oesophagus. 

The further course of the caeca is readily followed on the 
reconstructions (PI. IX, figs. 14, 15, 16) made from a typical 
specimen of Paragonimus kellicotli that was imbedded and 
sectioned without having been subject to pressure or other 
distortion. After the bifurcation of the oesophagus into the two 
lateral branches, these ascend rather rapidly toward the outer and 
upper margins of the body, and come to lie close under the 
vitellaria, but not widely separated from each other. This 
relation is apparent inasmuch as the body of Paragonimus 
kellicotti in a natural condition is not broad or flattened but 
attenuate and rounded. After its short transverse passage, each 
of the intestinal caeca passes backward toward the posterior 
extremity of the body, where they end blindly, one on each side, 
quite close together. The termination of each canal takes place at 
about the same level, although a slight variation occurs and 
sometimes one terminates a little sooner than the other. In lateral 
view the caeca show three large loops arching toward the dorsal 
surface. Between these, secondary loops appear. The three 
principal loops, while at first directed dorsally, ultimately curve 
with the body wall, and when viewed from above are seen to be 
turned also toward the median line. At these points the intestinal 
caeca approach each other very closely; in fact, the space between 
the most anteriorly located loops is very small. With this relation 
in mind it is evident that the digestive system is more extended than 
anticipated, and that each of the caeca if drawn out straight would 
reach nearly or fully twice the length of the entire body. There 
are also places where the intestine widens out considerably and 
shows distinct enlargements. In these places the diameter may 
become 0*68 mm., while towards the posterior extremity it is 
reduced to 0*225 mm. 

We are not sure how far it is possible to compare our 
description of the alimentary system in Paragonimus kellicotti with 
that of the other forms given by previous writers. We are not 
unconscious that those descriptions are possibly less detailed rather 
than actually different. What is certainly significant in explaining 
some variations is that much in previous accounts is taken from a 



126 


study of specimens flattened under pressure. Such specimens must 
be badly distorted, since this worm has a thick fleshy body and is 
reduced by the pressure to a mere fraction of its normal thickness. 
Nevertheless we feel that a comparison is worth while, and yields 
some minor evidence that is significant and helpful in the solution 
of our problem. On the whole it may be said that the anatomical 
study of the alimentary canal in Paragonimus westermanii and 
Paragonimus ringeri reveals some small but characteristic differences 
from that of Paragonimus kellicotti . The pharynx is distinctly 
smaller than in the latter. In Paragonimus westermanii it averages 
0'5 mm. long by 0*3 mm. broad, according to Kerbert; it is, then, 
not a spherical structure, as may be inferred both from these 
dimensions and from Kerbert’s figure. The pharynx of Paragonimus 
ringeri , according to Leuckart, is spherical and 0 3 mm. in 
diameter, but Kubo givfcs it as elongate, measuring 0*4 mm. by 
0*3 mm. The oesophagus is also shorter in both others than in 
Paragonimus kellicotti , being in Paragonimus westermanii 0*14 mm. 
long (Kerbert), and in Paragonimus ringeri 0*02 mm. long according 
to Leuckart. 

Kubo was the first to see that the regions of the two branches 
immediately following the median oesophagus are identical 
histologically with that canal, and should be counted a part of it 
and not of the intestinal caeca which continue them but are so 
different in structure. He gives a length of 0‘3 mm. for the 
undivided region and 0*2 mm. for the lateral branches. He 
comments on the sharp transition from the oesophageal to the 
intestinal region, and ascribes to the latter a diameter two to three 
times as great as we found in Paragonimus kellicotti . The peculiar 
condition shown in both these forms in that the intestinal caeca do 
not arise immediately from the end of the oesophagus, but some¬ 
what further distad from the branches of the median oesophageal 
canal, is no doubt a generic character. It is well illustrated by the 
figure of this region which Kubo gives. 

Some features of the intestinal caeca constitute possible 
differences. Kerbert says that in Paragonimus westermanii the 
intestinal caeca pass posteriad from the bifurcation of the 
oesophagus parallel with the surface of the body. They reach 
nearly to the posterior extremity of the body, and then end 



127 


blindly. He gives the length of the caeca at about 8 mm., or 
approximately the length of the whole body. This description is 
certainly at least incomplete, for the caeca in Paragonimus 
westermanii as he figures them do show some irregularities, and 
perhaps this condition varies with the age of the parasite in all 
species of this genus. But proceeding from the account of Kerbert, 
one may say confidently that such a simple relation does not exist 
in Paragonimus kellicotti or in Paragonimus ringeri. Leuckart 
shows, both in his description and by his sketches, that the 
intestinal caeca of Paragonimus ringeri are complicated by a 
number of loops and turns just as they are in Paragonimus 
kellicotti . Yet one point must be borne in mind, namely, that the 
latter parasite is the more attenuate. In it the intestinal caeca 
therefore do not lie very widely separated from each other, and 
where the major loops arch through the body towards the median 
dorsal line only a very small space intervenes. 

One may ask if the formation of the loops is the same in both 
species. According to Kubo the course of the intestinal caeca in 
Paragonimus ringeri is entirely irregular and very varied. This 
condition is shown in his figures, and may be a real difference 
between that species and Paragonimus kellicotti . We hesitate to 
accept this interpretation, since his figures are drawn from much 
flattened preparations, and in such the general symmetry of the 
caeca which we have described is so much modified that it can 
rarely, if ever, be seen. It is possible that an examination of 
undistorted specimens of Paragonimus ringeri will show some 
regularity and balance in the course of the intestinal caeca. For 
the present one must accept the statement that in Paragonimus 
ringeri the course of the intestinal caeca is irregular and 
asymmetrical. 

These relations between the species are expressed synoptically 
in the following table: 



P. westermanii 

P. ringeri 

P. kellicotti 

Size of Pharynx 

o*3 x o*5 mm. 

0*3 x 0*3 mm. or 

0*4 mm. 

0-44 x o*44 mm. 

Length of Oesophagus 

0*14 mm. 

0*2 or 0*3 mm. 

0*21 — 0*25 mm. 

Character of Intestinal 

Relatively simple, 

Looped irregularly (?) 

Looped symmetrically 

Caeca 

little longer than 
body (?) 

Twice length of 
body (?) 

Twice length of body 



128 


The Excretory System 

The excretory system in Paragonimus kellicotti is relatively 
simple. The excretory pore is of such size (0*05 mm. in diameter) 
that under favourable conditions its position may be determined 
with the unaided eye. It lies on the dorsal surface 0*2 to 0*225 mm. 
from the posterior extremity, and opens into a small canal about 
0 062 mm. in diameter and 0*225 mm. long. A cuticular layer lines 
the canal, surrounding which is a well-developed muscular layer. 
The canal passes directly forward, terminating abruptly approxi¬ 
mately midway between the dorsal and ventral margins of the large 
excretory sinus which lies in the middle axis of the body. The 
greatest dimension of this sinus is in a dorso-ventral direction, in 
which also it approaches the body surface very closely, coming 
within 0*15 mm. dorsally and 0*315 mm. ventrally behind the 
acetabulum. It extends anteriorly to within 0 45 mm. of the 
branching of the oesophagus, tapering to its termination here 
gradually, and finally disappearing nearer the dorsal than the 
ventral surface. As already stated, the sinus lies in the median 
axis, but in the region of the uterus, which is located on the left 
side somewhat anterior to the middle of the body; it shows a 
distinct bending to the right. Its median position is again resumed 
beyond the uterus. This bending is undoubtedly conditioned by 
the increase in size of the uterus as it becomes filled with eggs. 

The sinus has a distinct lining, as have also its main branches. 
These branches are very numerous, though along the sides of the 
sinus they are not conspicuous in the posterior region, but 
anteriorly in front of the acetabulum it is easy to see a number that 
are given off. These branches connect with the large stellate flame 
cells that are distributed throughout the body. We could not 
follow out the system to its ultimate details, but detected many 
flame cells in sections. 

The excretory system of Paragonimus westermanii consists also 
of the prominent elongated central reservoir and numerous lateral 
branches; in general a similar condition exists in Paragonimus 
ringeri and, as we have shown, in Paragonimus kellicotti . 
Considered more closely, however, there are here also certain minor 
differences. According to Kerbert the large central reservoir in 
Paragonimus westermanii is located in the posterior part of the 



129 


body, and opens to the exterior through a circular opening at the 
posterior pole. Kerbert also does not mention the presence of a 
short duct between the sinus proper and the excretory pore. This 
may be a minor error in observation, or due to methods of technic. 
But the appearance of the reservoir in Paragonimus westermanii is 
in most cases, according to Kerbert, that of an elongate or pear- 
shaped tube, or it may also be a spherical bladder. The latter 
observation was made upon fresh material. The excretory bladder 
in Paragonimus ringeri , according to Leuckart, is a very much 
elongated narrow sinus with its greatest dimension in the dorso- 
ventral direction. A short canal leads from the posterior extremity 
of the sinus to the excretory pore. This opening has a diameter of 
005 mm., and lies on the ventral surface. Our specimens of 
Paragonimus ringeri are slightly distorted, but seem to indicate 
this relation. In Paragonimus kellicotti the excretory pore appears 
on the dorsal surface in about the same relation with the posterior 
extremity. 

Kubo locates the pore at the posterior end. He also describes 
the canal system as originating from only two large main canals 
which empty one on each side into the posterior portion of the 
reservoir. Other authors have seen many such lateral canals 
emptying into the central reservoir, and our findings in Paragonimus 
kellicotti agree with them. 

The other relations are expressed in synoptic form as follows : 



P. westermanii 

P. ringeri 

P. kellicotti 

Location of 

Posterior pole 

Ventral, near posterior 

Dorsal, near posterior 

Excretory pore 


extremity, or at 
posterior pole (?) 

extremity 


The Reproductive System 

With the exception of the vitellaria,* the female reproductive 
organs of Paragonimus kellicotti extend only a little beyond the 
second quarter of the body. The genital pore lies about 0 078 to 
0*104 mm. behind the acetabulum, usually a little to one side of 
the median line. This is the opening of the genital cloaca. To 

• In this discussion we have retained the classic names for the organs although, since the 
appearance of Goldschmidt’s convincing demonstration, it can hardly be doubted that these 
designations are inappropriate and incorrect. 



130 


the right, and near the dorsal wall, is the ovary, while on the left, 
nearer the ventral surface, appears the highly coiled uterus. At 
about the level of the ovary, but in the median line, lies the shell 
gland, from which the proximal portion of the uterus emerges. 

The genital cloaca is a short flask-shaped structure only 0*2 to 
0’21 mm. long. It opens to the surface of the body through the 
genital pore, and receives the terminal ducts of both the male and 
the female reproductive systems. The wall of this structure 
consists of an inner homogeneous or granular layer, and an outer 
muscular wall which is composed essentially of circular fibres. The 
terminal portion of the vas deferens narrows to a small tube, the 
ductus ejaculatorius, which enters the genital cloaca at about the 
middle of the base. The metraterm, or terminal portion of the 
uterus, enters near the base, usually on the side opposite to the 
acetabulum. Cirrus and cirrus pouch are absent. 

Stiles records the genital pore as ‘median, right, or left, in 
specimens from hogs/ but in our specimens variance from the 
median location is slight, and where it is found, easily attributable 
to a slight distortion of the body surface. What Kubo calls the 
ductus communis genitalis evidently corresponds to what we have 
designated the genital cloaca. In our specimens it is certainly a 
distinct structure and not a common canal formed by the junction 
of the male and female ducts. It is very much smaller relatively 
than shown in his sketch, as is demonstrated by its length, which 
he gives as 0'4 mm. in Paragonimus ringeri . 

The male reproductive system of Paragonimus kellicotti 
consists of two testes that occupy the third quarter of the body, 
two vasa efferentia leading from them, and the single terminal 
vas deferens. The vasa efferentia unite near the dorsal margin of 
the excretory sinus to form the vas deferens. 

The testes lie one on each side of the body, and occupy nearly 
the entire space between the intestinal caeca and the excretory sinus 
in the posterior region of the body. The central portion of each 
testis is located approximately midway between the dorsal and 
ventral body surfaces. Their symmetrical arrangement is slightly 
disturbed, inasmuch as the right testis is usually a trifle posterior 
to the left testis. In this particular there is some variation, for in 
one specimen the right testis was found to lie anteriorly to the left. 



This variation will be discussed later. The form of the organs is 
noteworthy (Pl. VII, fig. 4, also PI. IX). Long slender lobes extend 
from the upper margin of the central mass; these are usually two 
in number. From the point of origin they arch upward and back¬ 
ward through the parenchymatous tissue. The terminal ends of 
the lobes are greatly enlarged, and frequently sub-divided into 
large rounded lobules. Other such lobes, three to four in number, 
extend from the ventral surface of the central mass. These 
connections are not as long as those given off from the dorsal 
margin, but their terminal ends are much more prominently 
enlarged, and almost always show two or more lobules heavier 
than those on the dorsal projections of the organ. 

The vasa efferentia, which are two in number, corresponding 
one to each of the testes, have a diameter of 0 026 to 0*046 mm. at 
their point of origin. They spring from the middle portion of the 
testes at about the same dorso-ventral level, although the right one 
is a little longer than the left. Each after its origin ascends 
gradually, and at the upper margin of the excretory sinus they lie 
parallel to each other. At this point the right one crosses to the 
left side of the body, and after both descend somewhat, they unite 
to form the vas deferens slightly to the left of the sinus at a level 
just below the shell gland and vertically above the genital pore. 

The vas deferens is at the start a relatively large duct, 0*062 to 
o* 1 mm. in diameter. It drops in general ventrad, arching first 
towards the anterior extremity. Keeping to the left side it 
approaches the acetabulum partly surrounded by the coils of the 
uterus. Then directing its course posteriad it circles close to the 
acetabulum towards the ventral surface, finally terminating in the 
genital cloaca. During its passage to the genital cloaca, the 
vas deferens shows a number of characteristic features. Just at the 
posterior margin of the acetabulum it suddenly narrows to a 
small tube, which becomes even smaller as the genital cloaca is 
approached. This portion of the vas deferens is heavily 
musculatured, no doubt functioning as the ductus ejaculatorius. 
Surrounding the vas deferens in the region where it suddenly 
narrows is a mass of glandular cells (PI. XI, fig. 21). These 
gradually disappear toward the genital cloaca and probably 
constitute the prostate gland. The inner cuticular lining of the 



132 


genital cloaca is continued into the ejaculatory duct, but farther 
on the nuclei of cells appear, although the lining retains its 
granular structure. A muscular layer consisting of circular fibres 
completes the wall of the vas deferens. 

The testes of Paragonitnus westermanii lie near the dorsal side 
of the body behind the transverse vitelline ducts (Kerbert). In 
structure they show five to six lobes. The right testis lies close 
behind the transverse vitelline ducts, while the left one is found 
nearer the posterior end of the body. For this reason it is possible 
to differentiate between an anterior right, and a posterior left testis 
which are distinct in Kerbert’s figure. It must not be forgotten 
that this was drawn from a much flattened specimen. The position 
and relation of the testes is different in Paragonitnus ringeri 
according to Leuckart. Here they lie, nearly symmetrically, well 
towards the posterior extremity of the body. They are not 
confined to the dorsal region, but occupy the greatest part of the 
space between the intestinal caeca and the excretory sinus. In the 
dorso-ventral dimension they have considerable extent. Comparing 
the figures given by Kerbert and by Leuckart, one sees a clear 
difference in form. The testes of Paragonimus westermanii are 
dense and the lobes more regular in form, while those of 
Paragonimus ringeri are diffuse and irregularly lobed. 

Kubo compares the testes of Paragonimus ringeri to an out¬ 
spread hand, and says they consist of four or five long lobes 
radiating from a common centre. Neither his description nor his 
figure will fit conditions in Paragonimus kcllicotti as we have found 
them, but they agree more nearly with Kerbert’s account of 
Paragonimus westermanii . Here again it is hard to say how much 
true conditions are modified by the distortion of flattened prepara¬ 
tions, but the testes in Paragonimus kellicotti are very much larger 
and both the central mass and the more numerous lobes are larger 
and heavier than the same structures as figured in the other species. 

Kerbert records that the vasa efferentia in Paragonimus 
westermanii pursue a dorsal course, arching over the transverse 
vitelline ducts; after several loops they approach the ventral surface 
and unite to form a common seminal vesicle which is continued 
into a short ductus ejaculatorius. The vasa efferentia in 
Paragonimus ringeri , according to Leuckart, pursue no such a 



133 


course, in fact neither one arches over the transverse vitelline 
ducts, while the left one drops gradually to the ventral surface 
without ascending dorsally. In addition, the vasa efferentia in 
Paragonimus westermanii are more slender, being o*oi to o*oi6mm. 
in diameter (Kerbert), while in Paragonimus ringert they are 0 045 
to 0*1 mm. in diameter (Leuckart). We do not feel sure to what 
extent these supposed differences, which are in fact rather minute, 
depend upon trivial errors in observation or description or upon 
different conditions of contraction in the body, and how far they 
indicate real variations in structure between these closely-allied 
species. 

As already stated, the vasa efferentia in Paragonimus ringeri. 
according to Leuckart, do not pursue a symmetrical course. It is 
important to examine this further. The right tube rises gradually 
toward the outer margin of the shell gland, crossing close under 
the transverse vitelline ducts, then drops almost perpendicularly, 
approaching at the same time the median line, and under the 
ventral margin of the shell gland unites with the vas efferens of 
the opposite side. This one pursues a much simpler course, for 
it does not approach the dorsal surface, but is directed downward 
toward the anterior extremity and median plane; it crosses the 
margin of the excretory sinus relatively far forward, and continuing 
its ventral and median direction is finally united with the other 
into the common duct. 

This portion of Leuckart’s description might be construed in 
either of two ways: (1) that the union of the vasa efferentia takes 
place ventrally to the excretory sinus, or (2) that it occurs dorsally 
to the sinus. The figure in the text illustrating this point shows 
the first relation, while the description might be understood as 
indicating either. Among the specimens of Paragonimus ringeri 
in the Ward collection there was one which had been broken just 
behind the genital pore. The anterior portion of this parasite was 
sectioned, and the relation of the vasa efferentia studied. In this 
specimen the ducts united dorsally to the excretory sinus. The 
vas deferens drops ventrally surrounded in part by the coils of the 
uterus. That there is opportunity for certain variation in the 
relation of the vasa efferentia, is readily understood from the fact 
that, with continued growth of the parasite, the uterus becomes 



! 34 

engorged with ova, finally pressing nearby structures out of their 
original relationships. 

The vitellaria of Paragonintus kellicotti are very extensively 
developed. Not only do they cover the parasite laterally, but also 
extend over the dorsal surface of the body, meeting both anteriorly 
and posteriorly, and leaving but a very narrow space in the median 
line which becomes broader just in front of the transverse vitelline 
ducts, finally to disappear entirely toward each extremity, although 
around the oral sucker there is also a free space. The ventral 
surface presents a relation very similar to the dorsal, except that 
on this surface the vitellaria do not meet near the anterior sucker, 
and correspondingly do not approach the median line so closely. 
The product of the vitelline glands is gathered up by many small 
ducts, which gradually unite to form two main trunks on each 
side, one arising in the anterior region and the other in the posterior 
region. These converge toward a point a short distance in front 
of the middle of the body, and unite here to form the large dorsally 
located transverse vitelline ducts. 

This distribution of the vitellaria stands in partial contrast with 
the condition in Paragonintus westermanii and Paragonintus ringeri. 
In these species, as described, a considerable space near the median 
dorsal and ventral line is not covered by vitellaria. In other words, 
the vitellaria on the dorsal surface of Paragonintus kellicotti 
approach more closely the median line than do those of 
Paragonintus westermanii and Paragonimus ringeri , and in other 
ways also appear to be more extensive in their development. The 
relation on the ventral surface is very similar. The vitellaria of 
Paragonimus kellicotti approach the median line more closely than 
those of Paragonimus westermanii and Paragonimus ringeri , but 
at the same time not so far as they do on the dorsal surface (PI. VII, 
fig. i). 

The vitelline reservoir in Paragonimus kellicotti is a pear-shaped 
structure arising at the point of union of the transverse vitelline 
ducts. These ducts become considerably narrower just before 
terminating in the vitelline reservoir. At the point where they 
unite the reservoir has its greatest width; dropping ventrally for a 
short distance and at the same time becoming narrower, it directs 
its course anteriad. Having reached a plane just below the shell 



!3S 


gland, it changes its course, and proceeds slightly upward until 
about the level of the junction of Laurer’s canal and the oviduct. 
Here it turns sharply to the right, and unites with the short canal 
formed by the union of these two ducts (PI. X, fig. 20). 

This condition closely typifies the relation in Paragotiimus 
westermanii , but not that described by Leuckart and Kubo for 
Paragotiimus ringeri. Here the two vitelline ducts unite to form 
a single canal, which drops ventrally a short distance, and then 
broadens out into a large flask-shaped reservoir 0 5 mm. long. 
From the median margin of the anterior extremity of this reservoir, 
and on the inner side of the unpaired canal, a small duct arises 
and passes dorsally to unite with the duct formed by the junction 
of the oviduct and Laurer’s canal. 

The ovary in Paragotiimus kellicotti lies on the right side, close 
to the dorsal wall of the body. Only a small portion extends 
down far enough to lie alongside of the excretory sinus. The 
transverse vitelline ducts bound this organ posteriorly (PI. IX, 
figs. 15,16). In relative size the ovary is about as large as a testis, 
but not so diffuse. It presents a more compact form since, even 
though lobed, the lobes are heavy and do not extend so far from 
the main body of the organ as do those of the testes. 

The oviduct is a short tube, at first relatively wide (o'36 mm.). 
It arises near the upper margin of the ovary, and from that portion 
which lies toward the median line. It soon narrows down to a 
very small tube, 0 018 mm. in diameter and about o'13 mm. long. 
Just after its origin on the ovary, the wall of the oviduct becomes 
heavily musculatured. This portion of the canal is the oocapt. 
The oviduct rises slightly toward the dorsal surface, but drops 
again towards the plane at which it left the ovary. Here it unites 
with a small duct, which in fact is the so-called Laurels canal. 
The wall of the oviduct consists of the cellular lining as is 
described for the male reproductive system, and the outer circular 
muscle layer (PI. X, fig. 19). 

Little can be said of these organs in a comparative way. The 
ovary lies in the same region of the body in all three species. We 
did not find any constant differences in its form or in the structure 
or relations of the oviduct in the different types. 

Near the beginning of Laurer’s canal is an expansion measuring 



0 057 mm. in diameter, and from this pocket there extends outward 
to the right a blind pouch, or small seminal receptacle, about 
o'195 mm. long and 0*052 mm. in diameter. Laurer*s canal makes 
its way from this pocket in a sinuous course towards the dorsal 
surface. While at the expanded region the canal is relatively 
large, it soon narrows down to a very small duct. It proceeds in 
a large loop directed posteriad, followed by another small loop in 
the opposite direction to a short vertical stretch directly above the 
shell gland which terminates on the dorsal surface of the body in 
the region of the transverse vitelline ducts. The expansion and 
the seminal receptacle, as well as the entire lower portion of 
Laurer’s canal, swarm with spermatozoa. The oviduct and 
Laurels canal unite to form a tube about 0*031 mm. in diameter. 
This extends only 0*045 mm - before it receives the vitelline duct, 
and then widens to form the ootype. The ootype discharges into 
the proximal end of the uterus. 

Stiles and Hassall (1900) have stated that a receptaculum 
seminis is lacking in the pig lung fluke. Their statement is not 
surprising in view of the real condition, which is shown at a glance 
in the figure representing the shell gland complex (PI. X, fig. 20). 
It is not possible, so far as we can determine, to detect such a 
structure in total preparations, but sections through this region 
demonstrate beyond question the existence of a true receptaculum 
seminis in its normal location. It has the form of a blind pouch 
opening into LaurePs canal near the inner end of the latter organ. 
At this point the canal itself is much expanded, and the receptacle 
can hardly be said to possess a neck or duct. Consequently the 
cavity of the pouch is in constant and open communication with the 
lumen of the canal, and spermatozoa circulate freely in the 
common space. The receptacle is small, measuring at most 
0*195 mm * length by 0 052 mm. in maximum width, so that its 
extreme tip does not even reach the border of the shell gland. 

The condition shown by the receptaculum seminis is of great 
interest from the standpoint of comparative anatomy. As is well 
known, there has been much discussion regarding the function and 
meaning of this organ. Most students regard it as a structure which 
is not of present functional value in any important way, at least 
among the trematodes, and certainly its variable character and 



*37 


occasional complete absence are strong arguments in favour of such 
a view. In some cases Laurer’s canal is reported to be lacking, and 
in others the receptaculum has been said to be wanting, as in the 
present instance. The actual presence of so insignificant a sac is 
only a theoretical correction of that statement. It may properly be 
designated a mere vestige of a structure about to disappear entirely. 
It is even possible, of course, that individual variation between 
different specimens is present to a sufficient extent to reduce it still 
further than the condition represented in the figure. So far we 
have found no evidence of its variation in size in Paragonintus 
kellicotti . 

According to Kerbert, the lower portion of Laurels canal in 
Paragonintus westermanii is provided with a seminal receptacle, 
which he figures somewhat larger than we find it in Paragonintus 
kellicotti. Leuckart, however, doubts the accuracy of this observa¬ 
tion, since he did not find such a structure in Paragonintus ringeri. 
A seminal receptacle certainly is present in Paragonintus kellicotti , 
and Kerbert not only records its presence in Paragonintus 
ivestertnanii , but also gives its measurements. If this structure is 
not present in Paragonintus ringeri , its absence may mark a 
characteristic difference in structure in the latter species. 
We are rather more inclined to believe that its presence can be 
demonstrated, and that if the three species differ at all in 
this respect, it will be found to be in the degree of development, 
or perhaps one should say, of reduction, which this organ 
manifests. Yet Kubo states that, despite zealous search, it was 
not possible for him to detect such a structure, and such a definite 
statement creates a strong presumption that the organ does not 
exist in Paragonintus ringeri. 

The shell gland as reported for Paragonintus ringeri by 
Leuckart is a large organ, lying a little to the right of the median 
dorsal region of the body. It is 0*5 mm. thick and about 1 mm. 
long. This organ is also well developed in Paragonintus kellicotti. 
It lies close to the dorsal wall, and is more or less oval in shape, 
although somewhat irregular in outline, and measures 1 mm. long, 
0 5 mm. thick, and 0 87 mm. broad. Kubo makes it slightly 
smaller, viz., o‘8 by o*6mm. Kerbert records approximately the 
same location in Paragonimus westermanii , although the shell gland 



>38 


is distinctly smaller, being 0’2 to 0*3 mm. long by 012 to 014mm. 
broad. The shell gland surrounds the proximal portion of the 
uterus, as well as the terminal portions of those ducts which go to 
form this part of the female reproductive system. 

The uterus in a fully-matured parasite is a condensed, closely 
coiled tube. As the ova accumulate the uterus becomes widely 
distended, so that the coil occupies nearly the entire lateral portion 
of this region of the parasite. The walls of the uterus arc made 
up of a relatively thin cellular layer, and a well-developed 
muscular coat consisting of circular fibres. Towards its terminal 
portion the uterus narrows down to form the metraterm. In this 
region the walls become heavily musculatured (PI. XI, fig. 22). 

The relation of the uterine coils in Paragonimus westermanii , 
as indicated by Kerbert’s sketch, is apparently simpler than is the 
condition in Paragonimus ringeri and Paragonimus kellicolti. In 
the former parasite the loops are open, and may be distinguished 
readily, but in the latter two forms they are close, and the entire 
organ presents the appearance of a solid mass. The age and 
stage of development must be a controlling factor in this condition, 
and very likely there is no constant difference here between the 
species. 

It is well known that among the Trematoda one finds at times 
an exact reversal in the usual position of the organs, chiefly of the 
reproductive system, so that the specimen is a mirror image of the 
usual relation. Such a condition has been designated amphitypy, 
and occurs in some species so frequently that it is impossible to 
say which is the normal and which the reversed situs genitalium. 
In Paragonimus such a reversal has been observed by several inves¬ 
tigators. Ward reported it for Paragonimus kellicotti , and we can 
confirm the record. In it the uterus lies on the left side, the right 
testis is anterior and slightly larger, the ovary is on the right side, 
and the bend in the excretory reservoir is to the right. This 
condition is not frequent in this species. Kubo found it in seven 
cases out of eighteen in Paragonimus ringeri. It is interesting to 
note that in the specimen figured by Looss (1914, p. 321) the organs 
are represented in what we regard as the reversed location, and 
not in the normal position. One figure published in Leuckart 
(1889) showed the same reversal. This has led to some confusion, 



139 

since the location of organs is stated by different writers in 
diametrically opposite terms. 

As one of us (Ward, 1908, p. 178) has noted previously, the 
correct measurement of ova is not a simple matter. All sorts of 
direct errors in counting micrometer spaces and in computing actual 
values are not only possible but, in actual practice, frequent. They 
are not easy to test or detect. Against the danger of computing 
averages from a small number of specimens it is hardly necessary 
to warn the student, although this error has been committed by 
some experienced men. A more insidious error is caused by the 
unconscious selection of the larger and more conspicuous specimens; 
in this way an investigator may raise the true average considerably 
and reduce the range of size through the elimination of the smaller 
specimens. Such a tendency can be detected if the full series of 
measurements is given, but can only be inferred with some hazard 
if the extreme and average measurements are the only data printed. 

The contrary difficulty, which is even more serious and more 
difficult to detect, will be introduced by that observer whose 
emphasis on the need of reporting every item he sees leads to his 
measuring and recording the size of absolutely every egg in a 
group. In any preparation one finds a considerable percentage of 
ova that are clearly abnormal. The shell is distorted by pressure 
or osmotic currents, which have made it over-large or over-wide; 
or in its process of manufacture by the parasite some interference 
with the normal course of events led to the moulding of an aberrant 
shell. The distorted form, or the atypical contents, demonstrate 
its unfitness for consideration in such measurements. To measure 
and record such an egg in the description of a species, unless the 
fact be given in connection with a note on the character of the 
individual egg, is usually to hamper, and certainly not to aid, 
the work of future students. 

Here we may quote again on this matter the views of Looss 
expressed in an article in this journal (1907, p. 149) which I cited 
previously in my discussion of this topic. He says : — 

‘ There exist, of course, among the immense number of ova in 
an individual worm always some which are either larger or smaller 
than the rest, or even evidently misshapen. In my opinion, it is 
of no use to record carefully the measurements of these eggs also. 



140 


For the description and definition of a species it is much more 
important to select for measurement those ova which appear to be 
normal, and to present the size and shape typical for the species. 
It may be added in passing that young worms with few ova in 
their uteri usually do not afford normally-shaped and normally- 
sized ova. 1 

Finally it does make a difference what is the exact source of 
the eggs measured. Those which are taken from the body of the 
worm near the beginning of the uterus, i.e., which are just formed, 
do not agree in size and proportions with those at the end of the 
uterus ready to be laid, or those which are collected after deposition 
by the worm. I have noted on several occasions a tendency of the 
student to measure ova from the first coils of the uterus, because 
there they are less crowded, and hence more easily seen and 
measured. With the passage of the eggs through the uterus they 
increase in size, probably by imbibition of uterine fluid, and the 
increase seems to be most marked in case the ovum enters upon its 
development during the intra-uterine period, so that when deposited 
the egg shell contains a more or less advanced embryo. 

Since one of us (Ward, 1908) has emphasized the significance 
of differences in the size of the eggs of Paragontmus as reported 
by various observers, especial attention was devoted to a study of 
these structures. An effort was made to get series of measure¬ 
ments under standard conditions, and to ascertain how far these 
varied from measurements previously given for eggs from similar 
sources. A start was made with Paragontmus kellicotti , and since 
it was evident at first thought that eggs taken from the body of the 
worm, or measured while still enclosed within it, might differ 
from those that had been laid in the normal manner, the first set 
of measurements was made from material that had been deposited 
naturally by the adult parasite. 

The mucous exudate obtained from the bronchi and from the 
worm cysts in infected hog lungs was brought carefully into 
glycerine jelly and mounted within asphaltum rings. These 
preparations contained large numbers of ova, and being protected 
from any pressure by the cover-glass, the ova were in a natural, 
undistorted and undamaged condition. 

Series of these eggs were measured, excluding only such 



specimens as were distinctly aberrant in form, or bore evidence of 
having suffered some mechanical injury. The maximum length 
obtained was 00875 mm. and the minimum o*0775mm., with an 
average of 0'o83 mm. The widest specimen measured 0*065 mm. 
and the narrowest 0 0525 mm., while the average width was 
0*0559 mm. 

The only other investigators who have measured these eggs 
under similar conditions are Stiles and Hassall, who state 
(1900, p. 603) that twenty-five eggs taken from cysts in the lungs 
of hogs varied in length from 0*096 mm. in maximum to 0*078 mm. 
in minimum, with an average of 0*0856mm., and in width from 
006 mm. in maximum to 0*048 mm. in minimum, averaging 
0*0532 mm. It will be noticed that so far as the length is 
concerned the minimum measurement of Stiles and Hassall is 
practically identical with our minimum, but the maximum is nearly 
ten micra larger. We were unable to find any eggs of that length 
in material mounted so as to remain free from pressure, and it is 
clear that they recorded very few since their average size is only 
slightly larger than our record. The width given in their records 
does not vary greatly from that we record. The slight differences 
of from 3 to 5 per cent, in the averages of length and width 
can not be regarded as of serious significance. These figures 
represent, undoubtedly, the approximate dimensions of the egg 
of Paragonimus kellicotti. 

In order to determine under similar conditions the size of the 
ova from the human lung fluke, slides were made in the same way, 
using the sputum obtained from an infected Korean. The material 
was not fresh as in the case of the hog parasite, but had been sent 
in formol from Chemulpo, Korea. A series of measurements from 
this material, after it had been treated exactly like the series of 
eggs from the sputum of the pig, gave the following values: — 
Length in maximum 0*097 mm., minimum 0*08 mm., with an 
average of 00872 mm.; breadth in maximum 0*055 mm.; minimum 
0*046 mm., with an average of 0*0506 mm. Some months later we 
obtained through the kindness of Dr. M. M. Null, of Seattle, 
Washington, sputum from the patient, a Korean also, in that city 
who had been found to be infected with the lung fluke (see Null, 
M.M., 1910). Four series of eggs from this material, after treat- 



142 


ment in the same manner as before, were measured separately, with 
the following results: — 



Length in mm. 

Width in mm. 


Maximum 

Minimum 

Average 

Maximum 

Minimum 

Average 

Scries (<j) ... 

0*0884 

0*0754 

0*0812 

0*0546 

0*0442 

0*0493 

Series (i b ). 

0*0884 

0*0780 

0*0822 

0*0520 

0*0468 

0*0499 

Series (r) . 

0*0858 

0*0780 

0*0806 

0*0546 

0*0442 

0*0496 

Series (d) . 

0*0858 

0*0780 

0*0813 

0 

N 

»/% 

O 

6 

0*0468 

0*0483 

General Average 



0*0812 



0*0492 


These series of eggs of the human lung fluke from different 
localities do not agree perfectly in measurements. This is 
especially noticeable in the length, which in the material from 
Korea had a much higher maximum, and consequently an average 
about 7 % greater. The width is almost identical, as the range 
agrees perfectly, and the average of the one differs only 2% from 
that of the other. Furthermore, it will be noted from the measure¬ 
ments cited that the range of variation in width is slight, being 
much less than the range of variation in length. It should be noted, 
as explained later, that the form of the egg is identical in the 
two cases. By reason of the extreme care used in obtaining these 
measurements, we believe they represent closely the true dimensions 
of the egg of this species and the probable range of size in normal 
undistorted eggs. 

There are many other observations on the eggs of the human 
lung fluke. In 1880 Baelz reported that the bodies he found in 
sputum measured 013 by 0 07 mm. This record may be rejected 
as unquestionably erroneous since it does not agree with any other 
report, and more especially since it was not cited three years later 
by the same author, who then gives the size of the ova as 0*08 to 
o' 1 by 0*05 mm. He gives no average size, but this range is close 
to that we obtained (0 08 to 0 097 by 0 05 mm.) for the Korean 
material. One other record is given for eggs taken fresh from 




! 43 

human sputum. Manson (1882) records the average size of these 
ova as 0*085 by 0*051 mm. 

Other records concern measurements made from preserved 
material or that of which the condition and method of handling 
is not stated. The most aberrant of these comes from Yamagiwa 
(1890), who measured in sections of the human brain and lungs ova 
which he attributed to the lung fluke, and found them in one case 
from 0 04 to 0*064 mm. by 0*024 to 0 04 mm., and in the other from 
0*049 to °’°6 mm. by 0*029 to 0*036 mm. These measurements are 
so extreme that they can not be accepted as belonging to the same 
species as that from which the other measurements were taken. On 
the other hand, they agree more closely with the eggs of 
Schistosoma japonicum , although smaller even for that species than 
the size given by others. Now Schistosoma japonicum has been 
discovered since Yamagiwa's work was published, and was not 
taken into account in his discussion. Without entering here upon 
any discussion of the other species, one may say confidently there is 
every reason for eliminating his record from the category of cases 
assignable to Paragonimus. 

Several other observations which undoubtedly concern the lung 
fluke remain for consideration. The record of Leuckart (1889) may 
have been made on material from the tiger or on some from the 
human host. He gives the length as varying from 0*08 to 0*i mrn., 
which agrees almost exactly with our figures, and states the width 
at 0 056 mm. which is barely larger than our recorded maximum 
width, 0*055 mm - These are distinctly larger than the figures 
given for ova of the tiger lung fluke, and hence probably were 
taken from specimens of the human lung parasite. Katsurada 
(1900) found the ova of the human lung fluke to vary from 0*0875 
to 0*1025 mm. in length with an average of 0*0935 mm., and from 
0*0525 to 0*0663 mm. in breadth with an average of 0*057 mm. 
Mackenzie (1904) records the length as 0*0855 to 0*0997 mm * with 
an average of 0*0913 mm., and the width from 0*048 to 0*069 mm., 
averaging 0*0552 mm. Both the extremes and the averages of these 
two records depart rather widely from our observations, and we 
are unable at this time to suggest any explanation for the 
discrepancy. 

Kubo did not devote particular attention to the egg, as he 



*44 

measured only ten specimens and gave but a very brief description. 
According to his records, the average length is 0*07 mm. and the 
average breadth 0 06 mm. These figures do not agree even 
approximately with those of any other investigator, and show a 
shell relatively much shorter and broader than anyone else has 
found. In fact, this average length is less than the minimum 
figures given in any previous paper, while at the same time the 
average breadth is greater than any other average, and equal to 
the maximum for this dimension given previously. 

These various results obtained from the measurements of eggs 
demonstrate wide discrepancies in technic or some inaccuracy in 
methods. That the eggs of Paragonimus really vary as much as 
these figures seem to indicate is hardly a tenable hypothesis. 
Looss, whose accuracy is unquestioned, says of these different 
records that he has personally found, in undistorted worms, eggs 
only from 0 077 to 0*081 mm. long and from 0 046 to 0*05 mm. 
broad, and accordingly the variant records undoubtedly belong to 
another species or to much distorted eggs in which the cover is 
easily forced off by pressure. We also have been unable to find 
any such variation in normal eggs as these figures given by different 
writers seem to indicate, even when the eggs are taken from adults 
in different hosts or from widely separated regions. Hence we are 
forced to assume the introduction of errors of some sort. 

In general it appears from these data that the eggs of 
Paragonimus kelticotti are broader and shorter than those of 
Paragonimus ringeri. These differences are not pronounced, and 
are certainly not sufficiently clear to allow of their use in the 
differentiation of species, as in extreme cases the sizes overlap. 
There is, however, a characteristic difference in form which may be 
seen on comparison of the non-operculated ends of the shell. In 
Paragonimus kellicotti that end tapers off rather sharply so as to 
produce the effect of a pointed extremity. In Paragonimus ringeri 
the corresponding end of the shell has a wider curve, due to a 
greater flare in the sides, which gives the egg a more pronounced 
elliptical outline. While this difference is trifling it is so distinct 
that once learned the student can readily pick out the species on 
comparison of the eggs. It was noticeable, for instance, that 
the two sets of ova from the human lung, discussed on pp. 141-2, 



differed somewhat in length from each other, but agreed perfectly 
in the outline of the non-operculate end of the shell, so that one 
of us measuring them at once recorded this correspondence in his 
notes before the data had been compared with other records or any 
attempt made to interpret them. 

Finally, it is noteworthy that the various records of size 
indicate much greater differences in length than in width, and there 
is a possibility that the eggs do vary widely in length while 
retaining a more uniform standard width. One could readily 
frame a hypothesis of egg formation which would agree with the 
known structure of the organs and the possible uniformity in width 
together with variation in length; but at present these supposed 
facts are too uncertain to be used in such a manner. 

The eggs of the Philippine forms were very carefully measured 
by Musgrave (1907, p. 35), using a Zeiss photomicrographic 
apparatus. In length they vary from 0*062 to 0'098 mm., with an 
average of 0*074 mm.; and in breadth from 0*047 to 0*063 mm. 
with an average of 0*057 mm. Compared with our own measure¬ 
ments it will be noted that the maximum length is in exact 
agreement, but the minimum and the average length are far too 
small. It may be, as Looss has suggested, that Musgrave has found 
and measured some eggs of Schistosoma due to an intercurrent 
infection. The elimination of these lower values will bring the 
' minimum and the average into agreement with our figures. But 
the width of the eggs as reported by Musgrave is open to the 
contrary objection. The minimum value agrees, but the maximum 
is far beyond what we found. This, of course, brings the average 
up above our record. It is only just to call attention to the fact 
that the figures of Musgrave for the width of the eggs agree fairly 
with those of the same dimension given by Katsurada and 
Mackenzie, although his values for the length of the eggs are very 
far below theirs. The evidence is inadequate for a final decision 
regarding the Philippine form. 

The eggs of the form found in the tiger measure 0*08 by 
0*045 mm., according to Kerbert; and Looss, who apparently 
re-measured the eggs of the same specimens sent to Leuckart, gives 
the dimensions as 0*077 to 0*o8i mm. long by 0*043 to 0*05 mm. 
broad. - These figures are practically identical, and as far as they 



146 


go indicate a slightly smaller egg than is found in the other two 
species. We did not have material for a re-examination and 
comparative study of these structures. The difference indicated, 
even if fully corroborated by a re-study of the material, is too 
slight to serve as a diagnostic basis for the species, since as in the 
case of the other species maximum values for Paragonimus 
westermanii exceed minimum values for Paragonimus ringeri. The 
same is true so far as length is concerned for Paragonimus 
kellicotti, but the egg of the latter species is always distinctly wider 
than the maximum value obtained for Paragonimus westermanii. 
The comment of Kerbert, not shown in his figures, that the eggs of 
Paragonimus westermanii are slightly flattened at the [both ?] 
poles, seems to furnish a slight difference in the form of the egg 
between this and the other species. 

GENERAL DISCUSSION 

The Genus Paragonimus 

Looss was the first to give an extended description of the genus, 
and his diagnosis slightly modified is reproduced by Stiles and 
Hassall (1900, p. 563). The description was so successfully 
written that even an extensive study of these three species has 
disclosed only minor corrections. Despite their distinctly insigni¬ 
ficant character it seems worth while to call attention to these 
characters. The pharynx, while well developed, manifests a 
tendency for a spherical form rather than an elongate, and even in 
the extreme case does not depart much from the spherical. In 
our opinion the oesophagus should not be designated as very short. 
It always appears in total preparations to be shorter than it 
actually is, since it rises obliquely towards the dorsal surface and 
thus is foreshortened in any longitudinal aspect, while in many 
specimens this inclination is so emphasized by contraction of the 
body that the branching of the caeca rests directly upon the 
pharynx and thus appears to originate from the latter without the 
intervention of any median unpaired region which is the oesophagus. 
The failure to see the oesophagus in total preparations is often 
responsible for the belief that such a region is non-existent when 
in lact it is well developed. In such cases it shows up 



conspicuously when the living worm stretches the anterior end, and 
becomes apparently obliterated when the worm assumes an average 
position or is contracted even moderately. In these instances the 
true form and relations of the oesophagus may be made out in 
lateral view, which one rarely gets of a trematode either naturally 
or in preparations, and may also be determined from a study of 
serial sections. From these it appears that the oesophagus suggests 
often the form of a letter S placed vertically, since it starts 
posteriad from a ventrally located pharynx, curves well to the 
front, and then turns again towards the posterior end to join the 
caeca. The beginning of these branches lies longitudinally so close 
to the pharynx that one does not see the vertical separation and 
the contracted or twisted oesophagus that joins them. 

Looss states that one testis lies obliquely behind the other, and 
this is hardly correct in any sense. As we have shown, the central 
mass of the one organ is located directly opposite that of the other, 
and the only difference is found in the lesser development of the 
anterior lobes in one, usually that on the right side. This is 
undoubtedly due to the fact that the total mass of the uterus which 
lies just anterior to that testis is greater than that of the ovary 
which is anterior to the other. When ovary and uterus are 
reversed, as has been reported to occur in exceptional cases, then 
the testes show the reverse of the usual development in that the 
anterior lobes of the left testis are less prominent than those of the 
right testis, and the organ appears to be located slightly posterior 
to that on the other side. The testes are in fact opposite each other 
and one appears slightly anterior to the other only because of the 
relative development of the anterior lobes. 

Both Looss, and Stiles and Hassall after him, emphasize the 
absence of a receptaculum seminis. As has been already shown in 
the discussion, this organ is present even though it be only poorly 
developed, and the proper form of statement would emphasize this 
fact. 

As noted above, all of these points are distinctly secondary, 
and the genus description remains substantially as outlined by 
Looss. On the other hand, one can hardly agree with him in 
regarding the genus as nearly related to the Fasciolinae, especially 
to Fasciolopsis, as he maintains it to be. Since we have just 



148 


learned through personal correspondence that Professor Odhner 
has in print a discussion of this matter, it seems best to omit here 
any elaboration of this topic. 

Stiles and Hassall (1900, p. 604) include in the genus two other 
species as follows: (A) Paragonimus rudis (Diesing, 1850) from the 
lung of a Brazilian otter; (B) Paragonimus comfactus (Cobbold, 
1859) from the lung of the Indian ichneumon. Neither of these has 
been reported since it was originally found, and of neither have 
we been able to secure material for comparison. 

Cuticular Spines 

It is important to point out more precisely the significance of our 
discovery concerning the cuticular spines. They constitute in our 
opinion a most convenient and accurate criterion for the distinction 
of species, and one which may well be applied to other genera 
among Trematoda to the advantage of the taxonomist on the one 
hand, and of the practitioner and pathologist on the other. To 
the latter, desirous of making a rapid and accurate diagnosis of a 
form which falls into his hands, such a definite feature will be of 
marked value. This is especially true since no complicated and 
time-consuming technic and no array of apparatus are demanded 
for the determination. It is sometimes possible to tear off with fine 
forceps a piece of the outer skin with some of the spines in situ; 
it is always possible to slice off freehand with a razor a thin layer 
of surface tissue containing them. Such a fragment, mounted 
roughly, gives a good view of the spines, and thus affords means 
for diagnosing the species once that the precise character of the 
spines has been determined for that species. This is pre-eminently 
a method for diagnosis from fresh material, since, as is well 
known, in many cases the spines are caducous in life and are easily 
lost also if the specimen lies some time in a preserving fluid. In 
fact, spines disappear or are overlooked so easily that many 
investigators have paid little attention to the statements regarding 
their presence or absence given by previous writers, knowing that 
the previously described specimens or their owm might easily have 
suffered the loss of these structures and yet be in good condition 
for the determination of other structural features. 



*49 


We should not neglect to say that in our experience these 
characters cannot be determined in specimens that have been 
reduced to series of sections. Several such series of Paragonimus 
have been loaned us by colleagues for the purpose of testing the 
character of these structures, and in no such case have we been able 
to determine anything concerning the precise form or the arrange¬ 
ment of the spines, even after long and painstaking effort. 
Undoubtedly such a series might contain a bit of cuticula, removed 
tangentially from a region where the spines were preserved well, 
that would give the desired evidence, but after our experience we 
are inclined to regard such an occurrence as exceptional, and to 
consider serial sections as unfitted to furnish the data desired 
regarding the cuticular spines. 

We have not been able to find record of a single species in 
which the precise form and distribution of these cuticular spines 
have been worked out. Yet they evidently possess advantages 
afforded by few structural features among the trematodes for the 
precise comparison of closely related forms. The great difficulty 
in comparing flukes results from the soft and variable form of most 
organs and the absence of hard parts. Descriptions are couched in 
terms that vary from individual to individual, and measurements 
in a given species range beyond the extremes of others, both among 
those truly similar and related and also among those that actually 
are more distant. The size of the suckers, which has been selected 
as one of the usual measurements in specific descriptions, varies 
considerably with the contraction of the specimen, and is moreover 
not always easy to determine, while it undoubtedly increases with 
the age of the specimen, although to what degree has not been fixed. 
Similar difficulties in the case of other organs contribute to make 
the description of the average trematode a recital of general, 
generic and even family characters rather than of specific features. 

Various investigators have sought to utilise the ova as characters 
for specific differentiation. The difficulties involved in placing 
dependence upon this factor are well exemplified by the discussion 
of the lung flukes in the preceding pages. Some important 
deductions have been made correctly on this basis, but the procedure 
is dangerous, as is any such argument based on a single factor. 
Furthermore, Looss has shown clearly that there are two species 



of Clonorchis in man, and yet the eggs are hardly distinguishable 
in size or form. 

The demonstration of another anatomical feature composed of 
material that cannot be altered by pressure, contraction, or other 
mechanical influence, and relatively constant in form among 
different individuals, furnishes a diagnostic element of distinct 
importance. When our attention was first drawn to the cuticular 
spines, and they seemed to furnish such an element, we felt the 
matter deserved the most careful study before any announcement 
was made. By the work reported in this paper one may fairly 
claim that a demonstration has been given for the essential 
similarity, even though not for the absolute identity, of these spines 
in different regions of the body of a single worm, and also on 
different individuals in the same host and in different hosts. We 
have been able to recognize a graduation in size and frequence 
characteristic of different parts of the body of the worm, and also 
a variation in size in different worms which may be due to the 
difference in host that harboured the specimens studied, but is more 
likely attributable to differences in age and growth of these 
structures in different individuals. In spite of the extreme 
differences in size and frequence, there is no approachment between 
the species, but rather added emphasis upon their real distinctness. 

In our opinion, these spines will furnish convenient and precise 
specific distinctions between still other species of Trematodes, and 
further studies to test this view are now in progress. It may be 
pointed out that this would be a most natural condition. Cuticular 
outgrowths, such as hairs, spines, scales, and other processes of 
varied form, have long been utilized by systematists in other groups 
as a means of identifying species, and have proved to be convenient 
and accurate characters for the description and differentiation of 
species. In animal groups which have been the object of long 
and intensive study, and in which taxonomy may justly be said to 
be more securely established than in groups of more recent study 
and hence less perfectly known, the use of such characters as specific 
criteria is generally approved. Even more than that, it may be 
said the species are distinguished on the basis of such differences 
when otherwise the structure, so far as worked out, is only known 
to be identical. Under these circumstances, we do not think we are 



venturing on dangerous or untried ground in maintaining the 
specific integrity of these three different forms of lung fluke, when 
the contention is supported by differences as clear and unmistakable 
as those we have demonstrated between the spines. The larger, 
more striking morphological differences, which some years back 
were regarded as specific in value, are now generally accepted as 
of generic or family rank. The proper evaluation of those items 
which are confessedly insignificant will yield a firm basis for the 
proper conception of species among the lower forms. Nowhere is 
there more need of such careful analysis of minor features than 
among parasitic worms, where great confusion reigris by virtue of 
the rapid and inexact treatment that has been accorded these forms 
in the past. Many investigators of the present day have broken 
away from that unfortunate tendency, and are analyzing the 
structure of such forms with great care. We hope that the factor 
to which we have called attention so prominently in this paper may 
prove to be of wider usefulness than in this case merely. Even 
should that not prove to be true, we are still confident that species 
of Paragonimus can be readily and accurately determined by 
means of the cuticular spines. Such a precise determination is of 
evident general interest, since only by it can be determined the 
range of territory infected by a definite form, the number of 
different species that threaten man in a given place, and ultimately 
the measures that must be taken to eliminate these parasites from 
the list of the enemies of man. 



152 


REFERENCES 


Fehleisen, F., and Cooper, C. M. (1910). Paragonimiasis or Parasitic Hemoptysis. Jour. 
Am. Med. Assn., LIV, pp. 697-699. 

Garrison, P. E., and Lxynes, R. (1909). The development of the Miraadium of Paragonimus 
under various Physical Conditions. Philippine Jour. Sci., B, IV, pp. 177-83. 

Hanson, H. (1911). Distoma pulmonale in Wisconsin. Johns Hopkins Hosp. Bull., XXII, 
pp. 112-4. 

Katsurada, F. (1900). Beitrag zur Kenntniss des Distomum toestermanii. Beitr. z. path. 
Anat. u. allg. Path., XXVIII, pp. 506-522. 

Krrbirt, C. (1881). Beitrag zur Kenntniss der Trematoden. Arch, fur mikr. Anat., XIX, 

pp . 5*9-578- 

Kubo, N. (1912). Morphologic des Distomum pulmonale. Centralbl. Bakt. u. Par., Abt. 1, 
Orig., LXV, pp. 115-138; 13 figg. 

Leuckart, Rud. (1889). Die Parasiten des Menschen und die von ihnen herriihrenden Krank- 
heiten, 2 Aufl., 1 Abt., 4 Lief., pp. 404-440. 

Looss, A. (1899). Weitere Beit rage zur Kenntniss der Trematoden-Fauna Aegyptens. 
Zool. Jahrb., Syst., 12: 521-784; 9 pi. 

-(1905). Von Wfirmcm und Arthropoden hervorgerufene Erkrankungen. Mense’s 

Handb. der Tropenkrankh., I, pp. 77-209; 2 pis. 

- (1907). Some Parasites in the Museum of the School of Tropical Medicine, Liverpool. 

Annals Trop. Med. and Par., I, pp. 123-152. 

- (1914). Wiirmer und die von ihnen hervorgerufenen Erkrankungen. Mense’s 

Handb. d. Tropenkrankheiten, II Aufl., II, pp. 311-516. 

Mackenzie, A. D. (1904). A case of parasitic hemoptysis or infection with the Distoma 
toestermanii. Jour. Am. Med. Assn., XLII, pp. 1133-1135. 

Manson, Patrick (1880). Further observations on Micro-Filariae, with Descriptions of 
New Species. Jour. Queckett Micr. Club, VI, pp. 138-139. 

- (1882). Distoma ringeri and parasitical haemoptysis. China Imperial Maritime 

Customs. Medical Reports, XXII, pp. 55-62 ; figs. 1-25. 

Musgrave, W. E. (1907). Paragonimiasis in the Philippine Islands. Philippine Jour. Sci., 
B, II, pp. 15-65 ; 11 pl». 

Nickerson, W. S. (1911)* Paragonimus in a Cat in Minneapolis. Science, n.s., pp. 33-271. 
Null, M. M. (1910). Tochil, or Endemic Hemoptysis. Northwest Medicine, II, pp. 364-366. 

Stiles, C. W., and Hamall, A. (1900). The Lung Fluke (. Paragonimus toestermanii) in Swine 
and its Relation to Parasitic Haemoptysis in Man. Ann. Rept. Bur. An. Ind., XVI, 
pp. 560-611 ; 2 pis. 

Ward, Henry B. (1894). Ueber das Vorkommen von Distoma toestermanii in den Vereinigten 
Staaten. Centr. f. Bakt. u. Par., XV, pp. 362-364. 

-(1903). Data for the Determination of Human Entozoa. Trans. Amer. Mic. Soc., 

XXIV, pp. 103-138 ; 4 pis. 

-- (1908). Data for the Determination of Human Entozoa II. Trans. Amer. Mic. Soc., 

XXVIII, pp. 177-202 ; 1 pi. 

Yamagiwa, K. (1890). Beitrag zur Aetiologie der Jackson’tchen Epilepsie. Arch. f. path. 
Anat. u. Phys., CXIX, pp. 447-460. 




* 5 + 


EXPLANATION OF PLATES VII-XI 

All figures are made from camera drawings of microscopical 

preparations, unless otherwise stated. 

Plate VII 

Fig. i. Paragotiimus kellicotti \ total preparation seen from the 
ventral surface. The vitellaria are represented on the 
left side of the worm, and omitted from the other side 
in order to show ovary, testis, vitelline ducts, and 
intestine normally obscured by them. The specimen had 
been stained, flattened under pressure, and mounted, 
x 7 5 diameters, a Egg from same specimen, x 300 
diameters. 

Fig. 2. Eggs of Paragotiimus kdlicotti showing ordinary varia¬ 
tions in form. These specimens were taken from mucus 
of lung, x 475. 

Fig. 3. Eggs of Paragotiimus kellicotti taken from uterus of 
parasite, x 475 diameters. 

Fig. 4. Reconstruction of a series of sagittal sections of 
Paragotiimus kellicotti showing the left half of the body 
in the posterior region as seen from the median sagittal 
plane. One can distinguish readily the marginal 
vitellaria, the dorsal shell gland with a bit of the main 
yolk duct or yolk reservoir, the uterus massed around 
the acetabulum, the irregularly-lobed testis with its 
vas efferens joining the vas deferens, and finally the 
extreme posterior tip of the intestinal caecum of that 
side. 








156 

Plate VIII 

Fig s . 5-11. Spines of Paragonimus kellicotti to demonstrate 
the substantial identity of these structures from different regions of 
the same specimen and pn parasites of the same species obtained 
from different hosts. 

Fig. 5. From dorsal surface behind oral sucker. 

Fig. 6. From centre of dorsum. 

Fig. 7. From dorsal surface near posterior extremity. 

Fig. 8. From ventral surface between oral and ventral suckers. 

Fig. 9. From region near oral sucker. 

Fig. 10. From region close to ventral sucker. 

Fig. 11. From specimen taken from lung of cat. 

Fig. 12. Spines from cuticula of Paragonimus ringeri from lung 
of man in Japan. 

Fig. 13. Spines from cuticula of type specimen of Paragonimus 
westermanii. 



Annals Trop. Med . & Parasitol.> Vol. IX 


PLATE VIII 



C. Tinling & Co., Ltd., Imp . 







158 


Plate IX 

Reconstruction of a specimen of Paragonimus kellicotti from 
the pig. The alimentary and reproductive systems alone are 
represented. The vitellaria are omitted and also the uterus, except 
in fig. 15, where it is outlined in part. The specimen was carefully 
preserved, and showed no evidence of distortion. It had not been 
flattened under pressure, as have most specimens drawn for illustra¬ 
tions in the texts. Consequently it shows more accurately the true 
form and relations of organs. 

Fig. 14. In lateral aspect, showing the right side of the body 
only. 

Fig. 15. In lateral aspect, showing the left side of the body only. 
Fig. 16. In dorsal aspect, showing both sides of the body. 



Annals 7rop. Med. & ParasitolVol. IX 


PLATE IX 



C. Tinting & 1 Co., Ltd., Imp . 






i6o 


Plate X 

Fig. 17. Cross section of intestinal wall, showing character of 
epithelial cells and subjacent tissue. 

Fig. 18. Cross section of body wall, showing cuticular spines 
in situ . The anterior end of the body lies to the right 
in the figure. 

Fig. 19. Section through ovary and oviduct, including also some 
cells of shell gland and the wall of the seminal vesicle. 

Fig. 20. Sagittal section supplemented from adjacent sections to 
show shell gland complex, including vitelline reservoir 
and common yolk duct, oviduct, seminal vesicle, Laurer's 
canal, ootype, shell gland, and beginning of uterus. 








Plate XI 

Fig. 21. Cross section of vas deferens near acetabulum. In 
this region the tube is filled with spermatozoa and 
surrounded by a loose mass of prostate gland cells. 

Fig. 22. Frontal section passing obliquely nearly through the 
length of the metraterm, which here contains a single 
egg. The layers of the wall and adjoining gland cells 
are clearly shown. 

Fig. 23. Cross section of oesophagus, showing layers in wall and 
associated salivary (?) gland cells. 



Annals Trop. Med. ParasitolVol. IX 


PLATE XI 



C. Ttitling dr 1 Co., Ltd., Imp. 





163 


PRELIMINARY NOTES ON THE 
MOSQUITOS OF KABINDA (LOMAMI), 
BELGIAN CONGO 


BY 

Dr. J. Schwetz 

(Received for publication July , 1914) 

Kabinda is a healthy station situated at an altitude of about 
2,800 feet. Malaria is known to exist, but is somewhat rare and of 
little importance. The habitations of the whites are grouped on 
the summits of several hills which are separated by ravines of 
varying depths. There are no swamps in the immediate 
neighbourhood of Kabinda, and mosquitos, in general, are 
relatively rare. They are, however, very sensitive to seasonal 
variations, and are more numerous at the beginning and end of the 
rainy season, when the rains are not excessive. 

At the time of my first visit to Kabinda I caught on myself, in 
the doctor's quarters, a specimen of Stegomyia fasciata, the presence 
of which species had already attracted my attention. Afterwards, 
in addition to several 4 ordinary' Culicine mosquitos, I captured a 
yellowish species (7 aeniorhynchus) and an Anopheline ( Myzontyia ). 
Thus from the beginning I observed that, in spite of the relative 
rarity of these flies at Kabinda, representatives of different groups 
were present, and that, therefore, an investigation would probably 
be of value. 

Returning later in order to pass some months at Kabinda, 
I settled down to the task of breeding these insects. There was no 
difficulty in obtaining the larvae, since it was only necessary for me, 
or a trained helper, to make a tour of the European or certain native 
compounds. Here would always be found the ‘ reservoirs containing 
stagnant water/ so strictly proscribed by malarial prophylaxy, 
namely, all kinds of receptacles containing rainwater or water 
intended for pigeons, poultry, ducks, etc. These receptacles, which 
are scarcely ever emptied, sometimes swarm with mosquito and 
Chironomid larvae. 



164 


Within a few months 1 bred a great number of mosquitos, 
several thousands, but before enumerating the tribes and important 
genera obtained (I am not sufficiently competent to determine with 
certainty the ‘ smaller ’• genera) I wish to make the following state¬ 
ments : — 

(1) I have not been able to breed certain species which, however, 
occur in the houses of the station. On the other hand, I have 
obtained in this way a mosquito which I have never seen in, or even 
near, the houses. Thus, the species of Taeniorhynchus are quite 
common in the houses, and I have seen them every evening in my 
own; nevertheless, I have not bred a single specimen. The same 
may be said for the members of the genus Mansonioides , which, 
though very rare, are also to be met with in the houses of Kabinda, 
and yet have never been bred. This probably means that the 
larvae of Taeniorhynchus and of Mansonioides require a special 
type of stagnant water, probably swamp water, and are not able to 
live in the more or less filthy water of the receptacles occurring near 
human habitations.* A few analogous facts which I have observed 
in other localities tend to strengthen this hypothesis. In the 
proximity of the station Mutombo- Mukulu there are numerous 
marshes, and mosquitos are very prevalent. 

During my first visit to this station, the mosquitos captured in 
my temporary habitation belonged almost exclusively to the genus 
Mansonioides. At the time of my second visit (the station had in 
the meantime been removed to a plateau a short distance further 
from the marsh), the mosquitos taken in the house were nearly all 
the yellowish Taeniorhynchus. During this second sojourn at 
Mutombo-Mukulu, however, I bred a certain number of mosquitos, 
and yet did not obtain a single specimen of Mansonioides or 
Taeniorhynchus. The same facts were observed at Samba 
(Kasonga Niembo). On the other hand, I have succeeded in 


# The immature stages of Mansonioides have been found by Dr. Ingram, in the Gold 
Coast, in borrow pits overgrown with a water plant (Pistia stratiotes). The larvae remain 
constantly below, attached, by means of their peculiarly modified siphon tubes, to the roots 
of the plant; the pupae, although not remaining continuously below, also attach themselves 
to the plant, by grasping the roots with their extremely elongate air-trumpets, and apparently 
pass considerable periods in this manner. The larvae of no African species of T aeniorbynebus 
have yet been discovered, but the life-histories of three or four American species have been 
worked out and are very similar to tho^e of Mansonioides. 


H. F. Carter 



raising a few examples of Megarhinus>* having found a few larvae 
in a receptacle, although these mosquitos are not seen in the 
proximity of habitations. This enormous mosquito, with the bent 
proboscis, does not occur in the houses, because it does not bite. 
But the fact that I have found larvae (four) of Megarhinus* in a 
small receptacle containing dirty water and that these larvae 
produced adults, shows that this mosquito has no need of special 
water or of any particular water-plant. 

(2) Generally speaking, the larvae of different species are not 
found in the same receptacle in any great variety. Those of a 
genus, or even of a species, not only predominate, but often are the 
only ones present at a given moment, and therefore the contents of 
a receptacle provide, as a rule, only a single species at a time. The 
same receptacle has furnished on one day specimens almost entirely 
of Culex pipiens,\ and eight days afterwards almost exclusively of 
Stegomyia fasciata. This is probably due merely to the chance of 
oviposition. 

(3) The Anophelines are relatively rare at Kabinda, both in the 
adult stage in and near houses, and in the larval stage, in artificial 
receptacles containing stagnant water. Among the thousands of 
mosquitos which have been bred, only a few dozen were Anophelines; 
these were obtained sometimes as rare examples among Culicines, 
and on one occasion as a pure ‘ culture 1 (associated Chironomid 
larvae having been removed). 

Among the Culicines, two important genera, Culex and 
Stegomyia , constituted 90 per cent, of those found. 

DETERMINATION AND CLASSIFICATION OF MOSQUITOS 
CAPTURED AND BRED AT KABINDA 

Not being a specialist, I have had and still have great difficulty 
in determining the species bred, all the more so since I have been 
unable to consult the special works on the subject. The exact 
identification is rendered still more difficult owing to the recent 
changes, wrought by Mr. Edwards, in the classification and 
terminology proposed by Prof. Theobald. As I wished above all 
to make this work of value, I resolved to avoid doubtful examples, 

# The tribe or sub-family Megar bin ini is intended, not the genus Megarhinus . H. F. C. 
t Probably C. iuttoni, Theob. H. F. C. 



i66 


and have, therefore, confined my determination to specimens which 
I recognised. I have sent the others for identification to Mr. F. W. 
Edwards (British Museum) and to Prof. R. Newstead and 
Mr. H. F. Carter (Liverpool School of Tropical Medicine), and 
have asked them at the same time to revise my determinations. I 
shall, therefore, be able at a further date to complete and correct 
this small preliminary study. 

The Culicinae (true mosquitos) are divided into four tribes— 
Megarhinini , Sabethini , Culicini , and Anophelini —and representa¬ 
tives of each have been found at Kabinda. 

(1) Megarhinini. In a receptacle containing water for the use 
of pigeons, I discovered, on one occasion, among some Culicine 
larvae, four enormous larvae which, from a distance, resembled 
small fish. Although I had never seen any so large, I immediately 
surmised, from their size, that they were probably the larvae of a 
Megarhine. Such an opportunity being so rare, I decided to 
preserve a larva and pupa; on the appearance of one of the latter 
a larva was killed, and when a second pupa developed one of these 
was also killed. The two remaining larvae hatched into two superb 
Megharines, a male and a female, of the same species, viz. : 
Toxorhynchites brevipalpis , Theob. The pupal stages of these 
two mosquitos were of considerable duration, and occupied five days 
in the case of the female, eight days in the male, whereas this period 
in other mosquitos is usually only one or two days. 

(2) Sabethini. These mosquitos were found several times in 
very small numbers among Culicines; twenty in all have been bred 
out, and two specimens were taken in the doctor's house. The 
characteristics of the larvae were not observed; all belonged to the 
same species— Eretmapodites chrysogaster , Graham. 

(3) Anophelini. As previously mentioned, the Anophelines 
are uncommon in this district; the few dozen examples raised from 
larvae, and the few rare specimens captured, belonged to the genus 
Myzomyia , Blanch. The species obtained at Kabinda and at all 
the other localities examined in this region—Mutombo-Mukulu, 
Samba, Labefu, Pania, etc.—proved to be M. costalis , Loew. 

(4) Culicini. This is the commonest tribe of mosquitos but the 
richest in genera and species, containing species of very variable 
size and colour. In consequence, it is the most difficult group to 



study, and the most intricate as regards classification. I have so 
far observed representatives of six different genera of the tribe, but 
have only been able to determine four with certainty. 

(a) Genus Culex , Linn.; C. tigripes , Grand.* Five examples of 
this species have been bred, the large larvae being associated with 
those of other species of Culex . C. pipiens group t—at least 50 per 
cent, of the mosquitos which I bred belonged to this group. 

(b) Genus Mansonioides , Theob. As previously mentioned, 
the larvae of these mosquitos have not been found, but I have 
captured a few adults of M. unifomits ,+ Theob., in the houses. 

( c ) Genus T aeniorhynchus % Arr. A score of adults were taken 
in my house, although none were bred. All the examples are of the 
same species, but show slight variations— T. aurites , Theob. § 

(< d ) Genus Stegomyia , Theob. After Culex , my breeding 
experiments have furnished me chiefly with members of this genus ; 
and almost all my specimens have been found, on very careful 
examination, to belong to the famous species, 5 . fasciata , 1 T Fab. 
5 . fasciata is very common here, and occurs both in and around the 
houses, chiefly towards evening , at which time, after sunset, they 
seem to be very active. The following curious fact may be suitably 
mentioned here. The mosquitos found in my house belonged, 
almost without exception, to the two genera Taeniorhynchus and 
Stegomyia . Now while the members of the genus Stegomyia 
became active and commenced to bite about sunset, it was almost 
exclusively those of Taeniorhynchus which annoyed me later, 
between seven and eight p.m.; before this they scarcely ever 
appeared. 

Besides the four more important genera dealt with above (and 
which are of interest in tropical medicine) I have bred numerous 
specimens of two other genera of Culicines. Since I am in doubt 

• Var. fusca, Theob. H. F. C. 

t Judging from the species present in the collection forwarded by Dr. Schwetz, it would 
seem that this term refers to C. duttoni , Theob., C. univittatus , Theob., and C. invidiosus, 
Theob. ; the true C. pipiens , Linn., was not represented. H. F. C. 

t Only a small number of specimens referable to this genus has been received from 
Dr. Schwetz, some of which may be Af. ajricanus , Theob. Owing to their poor condition, 
however, it is impossible to identify them with certainty. H. F. C. 

§ The examples forwarded proved to be T. cristatus , Theob., not T. aurites , Theob. 

H. F. C. 

H Other species of Stegomyia present in Dr. Schwetz* s collection were S. afrieatta. 
Theob. ; S. apieoargentea, Theob.; 5. smpsoni, Theob. ; S. potveri . Theob. II. F. C. 




i6H 


regarding these, I have sent the specimens for determination to 
persons more acquainted with the subject,* and shall refer to them 
in my next paper. A certain number of Chironomidae (other than 
Ceratopogon) and a few Psychodidae (other than Phlebotomus ) 
have also been bred, the larvae of these flies being found with those 
of the mosquitos. I shall mention these insects again later (as well 
as their larvae and pupae) when the specimens have been determined. 

My collection of mosquitos has been divided into four parts and 
sent to the School of Tropical Medicine, Brussels, the Museum of 
Tervueren, the British Museum and the Liverpool School of 
Tropical Medicine. 


• These specimens are referable to the species Culiciomyia nebulosa, Theob., and 
Ochlerotatus ( Protomacleaya) alboventralis , Theob. H. F. C. 



169 


ON THE PECULIAR MORPHOLOGICAL 
APPEARANCES OF A MALARIA 

PARASITE 


BY 

J. W. W. STEPHENS, M.D. (Cantab.) 

SIR ALFRED JONES PROFESSOR OF TROPICAL MEDICINE, THE UNIVERSITY OF LIVERPOOI 

{Received for publication 30 Nov., 1914) 

Plate XII 

In a paper entitled ‘A new Malaria parasite of man’ (1914), 
I described under the name Plasmodium tenue what I took to be a 
new species of malaria parasite. The peculiar morphological 
characters described were new to me, and I could find no illustra¬ 
tions or description of similar appearances. 

Since writing the above-mentioned paper I have received through 
the kindness of Dr. Le Fanu, W.A.M.S., from the Gold Coast, a 
blood slide in which he thought he had recognised the forms I had 
described as P. tenue. 

Dr Le Fanu had put this slide aside for further study, as he had 
noted before my description of P. tenue that there was something 
peculiar about the morphology of the parasites. The film was made 
from the blood of a native child that came for treatment. The 
infection is not a severe one: it is easy to find fields without any 
parasites. The forms in Dr. Le Fanu’s slide are, however, in some 
respects even more peculiar than the P. tenue forms. 

Firstly, I would point out that the blood cells present no evidence 
whatsoever of stretching or distortion, and the same peculiar forms 
of parasites occur in all parts of the film, thick and thin. 

Secondly, there are present large forms of quartan parasites 
which appear to me to be quite normal. In the slide it is possible 
to trace a transition from normal ring forms to those in which 
chromatin particles or strands without any protoplasm occur in the 
red cells. A better idea of these will be got from the accompanying 
plate than from any description however detailed. 

Three views seem to me possible as to the nature of these forms. 

(1) That they are a new species of parasite. 



170 


( 2 ) That they are degenerative, i.e., formed in the body under 
unknown conditions, and so perhaps analogous to the so-called 
quinine forms of parasites. 

(3) That they are artificial, i.c., formed outside the body under 
unknown conditions. 

The fact that among the parasites figured, forms are found in 
which chromatin alone without any protoplasm occurs in the red 
cell, is I think in favour of, but not decisive for, one of the latter 
two views rather than the first. The fact that normal quartan 
parasites are present is against these two latter views. It should 
be noted also that forms consisting of protoplasm alone, without 
chromatin, were not seen. Nor were pigment grains apart from 
parasites seen on the red cell. 

While this paper was passing through the press, Balfour and 
Wenyon (1914) published a paper entitled ‘The so-called 
Plasmodium tenue (Stephens).’ In it they express the opinion that 
I had ‘ not produced any evidence to prove that he [I] was not 
dealing with an amoeboid sub-tertian parasite.’ The fact that no 
observer (with perhaps one exception unknown to me when I wrote) 
since Laveran’s discovery of the malaria parasite in 1880, i.e., during 
a period of thirty-five years, had described such amoeboid forms of 
the malignant tertian parasite, appeared to me to be very strong 
evidence against this view. 

It is common knowledge that the malignant tertian parasite is to 
a certain extent amoeboid, but this is a very different condition from 
that I described in P. tenue. The authors quote for instance 
Ziemann, but a reference to his coloured plate will make it evident 
that he was not describing P. tenue forms. With regard to the 
possible exception I mentioned, I much regret that I had 
quite overlooked the existence of Dr. Balfour’s plate published in 
1908 and reproduced in the paper cited above. To Dr. Balfour 
certainly belongs the credit of the first description of markedly 
amoeboid forms of malaria parasites (other than the simple tertian), 
but a careful comparison of my plate illustrating P. tenue with that 
of Dr. Balfour will show that although there is a resemblance, it is 
not a close one. Now Ziemann’s plate was published in 1906, and 
if extraordinary amoeboid activity of the malignant tertian parasite 
was already recognised it seems hardly necessary for Dr. Balfour 
to have published his plate of ‘curious amoeboid forms’ in 1908. 



But to avoid controversial matter, let us now consider the 
substance of Balfour and Wenyon’s paper. They figure two 
parasites, the first of which (in a single film from West Africa) is 
almost certainly identical with P. tenue, and possibly, too, this is 
true of the second parasite (from Bagdad). In this latter case, several 
films were taken just before death and an hour previously; all 
contained the same forms (Dr. Wenyon in a private letter). As to 
the nature of these parasites the authors appear to be in doubt, for 
in one part of their paper they state that ‘ it may be possible to find 
for these variations some mechanical explanation,’ and in another 
that they are ‘young parasites’ (of P. falciparum ) ‘which are 
particularly amoeboid for some reason not clearly understood.’ 

Craig (1914), on the contrary, states that he ‘ is satisfied that 
Plasmodium tenue is an atypical form of Plasmodium vivax.' 

It is perhaps not out of place to mention here that Emin (1914) 
has described a new malaria parasite under the name Plasmodium 
vivax, var. minuta. 

When we have evidence as to what the nature of the forms I have 
described in this paper may be, and particularly whether they do 
or do not belong to the large quartan forms, we shall then probably 
be in a better position to estimate whether my view that P. tenue 
is a new species of malaria parasite is right or wrong. 

It is of interest to record the result of an examination of further 
films from the case of P. tenue. Two films (March, 1914) showed 
quartan, and in one of the films a single pigmented (presumably 
simple tertian) parasite was found with the pigment in the form of 
rods, in an enlarged cell showing Schuffner’s dots. Three films 
(June) showed quartan, and one (July), quartan. Young ring forms 
were not found in any of the films. 


REFERENCES 


Balfoct, A., and Wenyon, C. M. (1914). ‘ The So-called Plumodium tenu* (Stephen.).’ 

Jour, of Trop. Med. and Hyg., VoL XVII, p. 353. V 

Cxaig, C. F. (1914). 1 New varieties and .pecie. of Malaria plawnodia.’ Jour, of Para.it. 
(Urbana, Illinois), Vol. I, p. 85. 

Emin, A. (1914). ‘ Une vari£t6 nouvelle du parasite de Laveran.* Bull. Soc Path exot 

t. VII, p. 385. 

Stephens, J.W.W. (1914). * A New Malaria Parasite of Man.’ Proc. Roy. Soc., B. Vol 
LXXXVII, pp. 375-377 j and Annals of Trop. Med. and Parasit., Vol VIII 
pp. 119-122. ’ 



7 2 


EXPLANATION OF PLATE XII 

The figures were drawn with an Abbe camera lucida at a 
magnification of 2300 (approximately). 



Annals Trot>. Med. £* Parasitol., IW. /.V 


P/..-J7A A7/ 





ON SOME PREVIOUSLY UNDESCRIBED 
TABANIDAE FROM AFRICA 


BY 

HENRY F. CARTER, F.E.S. 

(LECTURER IN ENTOMOLOGY, LIVERPOOL SCHOOL OF TROI*ICAL MEDICINE) 

{Received for publication I February , 1915) 

Plate XIII 

The flies considered in this paper belong to the genera Tabanus 
and Haematopota of the sub-family Tabaninae; with one exception 
they were obtained either from the Transvaal or from the West 
Coast of Africa. Those from the Transvaal were all taken in the 
vicinity of Onderstepoort, near Pretoria, and formed part of a 
collection which also included the following species: Tabanus 
sericeiventrisy Loew, T. insignis , Loew, T. taeniola y P. de B., 
T. ditceniatuSy Macq., T. atrimanus, Loew, Haematopota scutellaris y 
Loew (?), Chrysops stigmaticaliSy Loew, and Diatomineura aethio - 
pica y Thun. 

The types and co-types of the various species and of the variety 
herein described are in the collection of the Liverpool School of 
Tropical Medicine. 


Genus TABANUS, Linn. 

Tabanus triquetrornatuSy n.sp. 

Medium-sized species with the dorsal surface of the thorax and 
abdomen chocolate-brown in colour; anterior half and posterior 
border of thorax, including whole of scutellum, pale greyish; hind 
margins of abdominal tergites with narrow pale borders expanding 
in the central region of the second, third, fourth and fifth visible 
segments into conspicuous greyish triangles and laterally into 
somewhat irregular pale areas. Eyes bare. Legs dark, the tibiae 
almost entirely white or creamy white. Wings faintly and uniformly 
tinged with brown. 



1 74 


Head (fig. l): Face, jowls and posterior region steel-grey, 
clothed with rather long white hairs; sub-callus (denuded) shining 
dark-brown. Front very narrow, from eight to nine times as long 
as the width at the base; yellowish-brown in colour, somewhat 
darker at the vertex, and clothed with short yellow and black 
hairs. Frontal callus shining, dark-brown, rectangular, reaching 
from eye to eye, and with a linear prolongation in the central 
line extending slightly further than the middle of the front. 



Fir,, i. (<i) Tabanus triquetrornatus, n.sp., front view of head, female, (x 13, about). 

(1 b ) Antenna of Tabanus triquetrornatus , n.sp., female, (x 13, about). 

Antennae (fig. ib) brownish-drab, darker, almost black, at the 
tips; first joint clothed with long white and short black hairs; 
second joint with a few white hairs ventrally and numerous minute 
black ones fringing the apical margins; third joint somewhat 
elongate with the dorsal process moderately pronounced and 
bearing a few short black hairs. Palpi creamy white, clothed with 
white hairs which, on the outer side of the terminal segment 
towards the apex, are intermingled with short black ones. 

Thorax : Anterior portion of dorsum, as far as the transverse 
suture, ashy-grey, clothed with yellow and black hairs, the former 
apparently* more numerous laterally; remaining portion of dorsum 


The dorsal surface of the thorax is partially denuded in this specimen. 



i75 


(except area immediately in front of scutellum, basal half of post- 
alar calli and scutellum , all of which are yellowish-grey) of a 
chocolate-brown colour; humeral and post-alar calli bearing long 
white hairs. Pleurae steel-grey, slightly darker anteriorly with 
whitish hairs. 

Abdomen : Dorsal surface chocolate-brown, clothed with short 
black hairs; distal margins of segments, triangular areas and lateral 
expansions light grey, with yellowish hairs; pale triangles on 
second, third and fourth segments large, their apices reaching 
almost to the basal margins of the segments on which they are 
situated, that on the fifth segment considerably smaller and only 
half the height of the segment. Venter , two basal segments entirely 
grey, clothed with white hairs; remaining segments chocolate-brown 
with pale grey hind margins expanding somewhat laterally, clothed 
with white hairs interspersed on the darker areas, with black ones 
which become longer and more numerous on the last two segments. 

Wings : Faintly brownish; veins and stigma pale reddish- 
brown. Squamae chocolate-brown. Halteres , stems and knobs 
brown. 

Legs: Coxae grey, clothed with white hairs. Femora black, 
more or less pruinose ventrally and laterally, especially those of the 
hind legs; dothed with black hairs which are intermingled with, 
or replaced by, white hairs on the pruinose areas. Tibiae, except 
extreme apices, white or yellowish-white clothed with white hairs. 
Apices of tibiae and whole of tarsi black, with short black hairs. 

Lengthy 12 mm.; length of wing \o' 2 $ mm., width of head 
4 mm.; width of front at vertex o’5 mm. 

Habitat: Calabar, Southern Nigeria; Dr. T. B. Adam, 1913; 
one female. 

This species is evidently related to Tabanus argenteus % Surcouf. 
It is easily distinguished, however, by its general paler coloration, 
paler wings, and by its abdominal ornamentation. 

Tabanus fuscipesy Ric., var. oculipilus n. var. 

Closely resembling the typical form, but with the eyes more 
thickly covered with longer hairs, the femora distinctly paler and 
the basal portion of the third joint of the antenna somewhat shorter 
and broader. The femora are yellowish, only the basal third of 



i 7 6 


the front pair and the extreme bases of the middle and hind pairs 
being dark. The recurrent veinlet from the upper branch of the 
cubital fork is usually considerably longer and more conspicuous 
than in the typical form. 



Fig. 2. ( a ) Antenna of Tabantis fuscipes var. oculipilus , n.var., female, (x 35, about). 

( 3 ) Antenna of Tabanus fuscipes, Ric., female, (x 35, about). 

Length , 11*25 to 13*25 mm.; length of wing, 875 to 10*25 mm - 

Habitat : Onderstepoort, Transvaal; type female, 2.2.1913, 
Mr. G. A. H. Bedford; thirteen females, presented by Mr. F. V. 
Theobald, taken in the same district during January, February and 
March, 1911. 

Both Tabanus fuscipes , Ric., and its near relation T . ditceniatus , 
Macq., belong to the sub-genus Atylotus , Osten-Sacken, all the 
members of which possess pubescent eyes. In these two species, 
however, the hairs are almost microscopical, whereas in the new 
variety the pubescence of the eyes is very marked and clearly 
visible under a pocket lens (x 15). Taking into consideration also 
the other differences mentioned it would seem that the form in 
question is worthy of varietal rank. T. taeniatus f Macq., which 
also occurs in South Africa and possesses hairy eyes, might perhaps 
be confused with this insect but may be distinguished, inter alia , 
by the presence of two sharply defined whitish, admedian, stripes 
on the thorax, by the abdominal stripes being white instead of 
yellowish, and by the legs and antennae being darker. 


Genus HAEMATOPOTA, Mcig. 

Haematopota transvaalensis , n. sp. 

(Plate XIII, fig. 3) 

Medium-sized blackish species with conspicuous, pale, more or 
less circular spots on each abdominal tergite, grey, clearly-marked 
wings and dark legs—the front tibiae strongly incrassate. 



i ?7 


Head (fig. 3) : Face , jowls and posterior region whitish, clothed 
with long white hairs; upper part of face with a row of four 
velvety-black spots extending horizontally just below the bases of 
the antennae; each of the inner spots is rounded and is situated 
immediately below one of the antennae, while each outer spot is 
more or less triangular with its base touching the eye; face above 
the line of spots yellowish-brown with numerous, irregularly 
distributed, small black spots. Front rather dark mouse-grey, 
except the extreme lateral margins and round the lateral frontal 
spots, where it is much paler and almost white; vertex yellowish- 
brown, but with a short, narrow, greyish stripe in the middle line 



Fig. 3. Haematopota transvaalensis , n.sp., side view of head, female, (x 18, about). 

extending to the median frontal spot; front clothed with black and 
white hairs, the latter present on the paler areas. Lateral frontal 
spots large, rounded, touching the eyes; median spot small and 
inconspicuous. Frontal callus shining, olive-brown, of moderate 
depth and extending from eye to eye; spot below callus black, or 
nearly so, and conspicuous. Antennae (fig. 3) comparatively long 
and slender; first joint cylindrical, dark greyish-brown, clothed 
with short black hairs; second joint and extreme base of third, 
yellowish-brown, the former clothed with minute black hairs; third 
joint, except extreme base, very dark brown, terminal annuli almost 



i 7 * 


black, the last annulus about the same length as the two preceding 
together. Palpi grey; terminal segment clothed with short black • 
hairs on outer side, replaced towards the base by a few rather long 
whitish hairs. 

Thorax \ Dorsal surface dark brown with three longitudinal 
light grey or whitish stripes; median stripe narrow, more clearly 
defined anteriorly, extending from front margin to scutellum 
(excepting a short distance in the centre where it is obliterated) and 
dilating somewhat posteriorly; lateral, admedian, stripes broader, 
terminating just above the inner extremities of the transverse suture, 
below each of which is a conspicuous, more or less triangular, white 
spot; side of dorsum and humeral calli darker grey, crescentic 
marks on hind margin sharply defined, whitish, each with the base 
of an ill-defined, moderately large, smoke-grey triangular area 
immediately above its outer extremity Scutellum dark brown; the 
front margin yellowish-brown in the centre merging into light grey 
laterally, sides and broad median stripe, which extends to hind 
margin, grey. Dorsum sparsely clothed with glistening white 
hairs, humeral calli with long white hairs, post-alar calli with white 
and black hairs intermingled, and scutellum with hairs similar to 
those of the main portion of dorsum. Pleurae light grey, clothed 
chiefly with white hairs. 

Abdomen : Dorsum dark brown, the lateral borders of the 
segments grey and the hind margins narrowly creamy-white; each 
tergite with a pair of comparatively large, yellowish to grey, 
admedian spots, and the third to sixth tergites each with a narrow 
grey median line; yellowish-grey hind margin of second tergite 
produced into a forwardly directed yellowish triangle. Dorsum 
clothed with black or brown hairs on the dark areas and with 
glistening white hairs on the remaining portions. Venter steel grey, 
with hind margins of segments paler, clothed with glistening white 
hairs; last segment darker with some erect black hairs. 

Wings : Grey with rather fine pale markings as shown in 
Plate XIII, fig. 3; rosettes well-defined, apical sinuous mark 
single, extending from costa in first submarginal cell (where it is 
expanded) to wing margin at a point just below the lower branch 
of the third longitudinal vein, distal angles of third and fifth 
posterior cells, each with a large pale blotch. Stigma conspicuous, 



179 


dark brown with the proximal third paler; veins brown. Squamae 
pale, tialteresy stalks cream-coloured, knobs dark brown. 

Legs : Coxae dark grey (upper half of front pair paler), clothed 
with whitish hairs. Femora mostly pruinose, clothed with white 
and black hairs, the former preponderating except on the front 
femora where the black hairs are more numerous (especially at the 
apices) and rather longer. Front tibiae black, distinctly swollen, 
with a conspicuous creamy-white ring at the base, middle and hind 
tibiae blackish-brown, each with two cream-coloured rings, hind 
pair slightly dilated; clothed with black hairs except on the pale 
bands where white or creamy-white hairs are present—on the hind 
tibiae the pale hairs extend, for some distance, on to the dark space 
between the rings. Tarsi black or dark brown, the first tarsal 
segments of the middle and hind pairs of legs mostly cream-coloured. 

Lengthy io mm.; length of wing, 9 mm.; width of head, 3*5 mm.; 
width of front at vertex, ri mm. 

Habitat : Onderstepoort, Transvaal; one female presented by 
Mr. F. V. Theobald, 24.3.1911, and one other female by 
Mr. G. A. H. Bedford, 9.12.1912. 

H . transvaalensis does not seem to resemble closely any of the 
species described by Loew in his work on South African Diptera, 
but is apparently related to Haematopota masculosifaciesy Aust. 
From this, however, it may be distinguished by its larger size, 
paler scutellum, differences in wing markings ( inter aliay the apical 
sinuous mark), and in its abdominal ornamentation. 

Haematopota theobaldiy n. sp. 

(Plate XIII, fig. 4) 

Medium-sized chocolate-brown coloured species with reddish- 
brown wings; dorsum of thorax paler with three longitudinal smoke- 
grey stripes, dorsum of abdomen darker with a conspicuous ashy- 
grey median stripe and traces of spots on, at least, the apical 
segments. First antennal segment much swollen and shining black. 
Legs dark, the front tibiae incrassate with one pale ring, mid and 
hind tibiae with two pale rings. 

Head (fig. 4): Face, jowls, and basioccipital region light grey, 
clothed with whitish hairs; upper part of face with a transverse row 



i8o 

of black spots, as in the preceding species, but less distinct, the 
inner ones, below the antennae, reduced to mere dots, the outer ones 
narrowly separated from the eyes. Front , clothed with glistening 
cream-coloured hairs, dark mouse-grey in the centre, light grey at 
the extreme lateral margins and round the lateral frontal spots; 



Fio. 4. Hatmatopota tbeobaldi , n.*p., side view of head, female. (x 18, about). 

vertex with a large dark brown blotch, which becomes somewhat 
yellowish-brown at the margins, divided in the central line by a 
narrow grey stripe reaching to the median frontal spot. Lateral 
frontal spots, more or less circular, not in contact with the eyes; 
median frontal spot much reduced and inconspicuous. Frontal 
callus shining black, of moderate depth and extending from eye 
to eye; upper margin slightly produced in the centre, lower margin 
concave for some distance on each side of the middle line; spot below 
callus velvety black. Antennae (fig. 4) dark, moderately long; first 
joint strongly incrassated and shining black (greyish pollinose at 
base and on inner surfaces), clothed with sparse black hairs; second 
joint dull reddish-brown, upper angle somewhat produced, clothed 
with black hairs; third joint dark brown, slightly paler at extreme 
base, with the terminal annuli black—last annulus not quite so long 
as the two preceding taken together. Palpi dark grey; first 
segment clothed with white hairs, second segment clothed on the 
outer side with short black hairs interspersed with rather longer 
white ones towards the base. 


Thorax : Dorsum chocolate-brown with three longitudinal, rather 
dark smoke-grey lines; median stripe narrow, paler at extreme 
anterior end, and extending to the scutellum; admedian stripes 
shorter, widening and becoming contiguous with median stripe 
anteriorly, terminating immediately behind the transverse suture as 
a more or less triangular spot; usual crescentic marks on the hind 
margin inconspicuous, sides of dorsum, including humeral calli, 
grey. Dorsum sparsely clothed with glistening yellow hairs; humeral 
calli with long whitish hairs. Scutellum chocolate-brown with hind 
margins and central region paler. Pleurae pale mouse-grey clothed 
with white hairs. 

Abdomen : Dorsally rather darker than thorax with a relatively 
broad ashy-grey median stripe, and paired, more or less circular, 
similarly coloured, ill-defined, spots on the second to fifth segments; 
hind margins of segments, and part of front margin of second 
segment, narrowly ashy-grey, and extreme lateral margins of tergites 
one to four, light grey, the pale area becoming gradually smaller 
towards the fourth segment. Hairs clothing dorsum similar to 
those on thorax, but interspersed with black ones, especially on the 
darker areas and on the apical segment; lateral pale markings clothed 
with white hairs. Venter pale mouse-grey anteriorly and laterally, 
gradually merging into chocolate-brown towards the apex, hind 
margins with very narrow pale borders; clothed with short yellowish 
hairs proximally, with black hairs distally, and with hairs of either 
colour on the central segments. 

Wings : Reddish-brown with rather coarse pale markings 
arranged as shown in Plate XIII, fig. 4. Stigma y elongate, dark 
reddish-brown, the basal third paler, yellowish-brown. Squamae 
whitish. Halteres with cream-coloured stems and dark brown 
knobs, the latter divided in the middle line by a fine creamy stripe. 

Legs : Coxae dark grey; those of the fore legs clothed with short 
black hairs and some longer white ones towards the base, those of 
the middle and hind legs clothed with short whitish hairs. Femora 
dark brown, paler and greyish ventrally, clothed both with black 
and white hairs—the latter situated chiefly on the pale areas. 
Tibiae dark brown; front pair almost black, strongly incrassated 
and with a narrow cream-coloured band at the base; middle and 
hind pairs with two rather inconspicuous pale bands; tibiae, except 



pale bands, clothed with black hairs, the bands with yellowish-white 
hairs. Front tarsi black, clothed with black hairs; middle and 
hind tarsi dark brown, the first segment of each mostly cream- 
coloured, and remaining segments of hind pair narrowly pale (cream- 
buff) at extreme base. 

Length , 875-10 mm.; length of wing 74-87 mm.; width of 
head 2 6-2'g mm.; width of front at vertex *9-105 mm. 

Habitat: Onderstepoort, Transvaal; three females presented 
by Mr. F. V. Theobald (to whom I have much pleasure in 
dedicating this species), 1911, and one female by Mr. G. A. H. 
Bedford, 1913. 

Mr. E. E. Austen, of the British Museum, has kindly examined 
one of the above specimens, and states that ‘in some respects it 
appears to resemble Hacmatopota bistrigata , Loew, but is dis¬ 
tinguishable at once from that species by inter alia the strongly 
incrassated front tibiae.* 

Haematopota pinguicornis, n. sp. 

Small, dark, strikingly marked species; dorsal surface of thorax 
dark brown with a very broad, pale, median, longitudinal area on 
the anterior portion and a light coloured posterior margin; dorsum 
of abdomen dark brown with narrow greyish hind margins to the 
segments. Antennae dark brown, the first joint strongly incrassated 
and shining, the third joint short and flattened from side to side. 
Wings sepia with very conspicuous pale markings. Legs dark with 
pale bands on the tibiae, front pair with one pale ring, middle and 
hind pairs each with two pale rings—the hind tibiae somewhat 
dilated. 

Head (fig. 5): Face y jowls and posterior region, grey, clothed 
with white hairs interspersed with a few short black ones; face with 
a conspicuous black transverse line extending from eye to eye, and 
passing immediately below the bases of the antennae, above this line 
the face, except the narrow whitish borders to the eyes, is yellowish- 
brown in colour becoming darker towards the antennae and margins. 
Front dull chocolate-brown with light grey lateral margins which 
extend inwards to enclose, or partially enclose, the lateral frontal 
spots and diagonally downwards, from the lateral comers of the 
vertex, to encircle the median frontal spot; vertex, between the two 



diagonally projecting grey stripes, dark brown but with a fine grey 
median stripe extending to the median frontal spot. All three 
frontal spots distinct, the median spot rather small and circular, 
the lateral spots well-developed and more or less quadrate. Front 
somewhat denuded, apparently clothed with black and white hairs, 
the white hairs being limited to the paler areas. Frontal callus 
shining, dark brown or black, of moderate depth and with a distinct 
depression in the middle line; upper margin almost straight and 
narrowly separated from the eye at each end, lower margin slightly 
convex, its lateral extremities in contact with the eyes. Spot below 
callus well-defined, dark brown, quadrate.. Antennae (fig. 5) dark, 
clothed on the hair-bearing portions with black hairs; first joint 



Fig. 5. Haematopota pinguicornis, n.sp., side view of head, 
female, (x 1 8 , about). 

much swollen, shining, dark brown but paler (yellowish-brown) 
towards the base and on the inner surfaces; third joint short, rather 
longer than the first and second joints taken together, with the 
expanded portion much compressed laterally and somewhat abruptly 
constricted basally; annuli fairly distinct, the last annulus about the 
same length as the two preceding together. Palpi grey; first 
segment darker and somewhat brownish, clothed chiefly with long 
white hairs, a few black ones present towards the apex; second 
segment rather narrow, clothed on the outer side with numerous 
relatively long black hairs, interspersed with a few white hairs. 

Thorax : Pale median area on dorsum extending longitudinally 
from anterior margin of thorax to slightly beyond level of transverse 



suture, its outer margins (corresponding to the usual admedian 
stripes) sharply defined, light grey, gradually darkening and 
becoming pale brownish in the central region, in which the median 
stripe may be traced as a narrow slightly paler line; light grey outer 
margins, immediately above inner lateral extremities of transverse 
suture, each curving inwards and uniting, near the middle line, with 
the inner edge of the light grey spot, situated below the inner lateral 
extremity of the transverse suture; crescentic marks on posterior 
margin large, light grey; humeral calli grey with white and black 
hairs intermingled, the white preponderating; sides each with an 
oval grey area, immediately behind humeral callus extending back¬ 
wards as a moderately broad line to outer extremity of post-alar 
callus where, after a short interruption, it curves inwards and 
upwards forming a somewhat conspicuous spot. Scutellum dark 
brown, paler towards the upper margin. Dorsum clothed with both 
black and white hairs, their distribution corresponding more or less 
to the darker or lighter markings. Pleurae dark grey, clothed with 
white hairs. 

Abdomen: Dorsum dark chocolate-brown, clothed with short black 
hairs, the hind and lateral margins of the tergites grey or yellowish- 
grey, clothed with white hairs. Venter , first segment greyish-brown, 
others with colouring and ornamentation similar to that on the 
tergites; clothed with short black hairs on the dark portions and 
glistening whitish hairs on the narrow grey margins, last ventral 
scute with numerous long black hairs. 

Wings (Plate XIII, fig. l) : Dark with sharply defined coarse 
pale markings; apical sinuous mark strongly bifurcate, both branches 
broad, the outer ramus situated close to the wing margin; posterior 
cells, except fourth, and axillary cell with conspicuous pale blotches 
in their distal marginal angles, fourth posterior cell with a small 
indistinct pale mark in its distal angle; other markings as in the 
figure. Stigma somewhat elongate, very dark brown, conspicuous, 
with a dark rectangular blotch below extending nearly to upper 
margin of discal cell. Squamae white. Halteres , stalks cream- 
coloured, knobs dark brown. 

Legs : Coxae dark brown, clothed with black hairs, basal half 
of front pair grey with long white hairs intermingled with the black 
ones. Femora dark brown, almost black, clothed with black hairs, 



middle and hind femora rather paler ventrally and laterally, the 
hind pair slightly thickened. Tibiae, front pair black, each with 
a narrow cream-coloured band near the base, middle and hind pairs 
dark brown with two pale rings; whole of tibiae, except on the pale 
bands, clothed with black hairs, the pale bands with white hairs. 
Tarsi, front pair black, middle and hind pairs dark brown, each 
with the first segment mostly cream-coloured. 

Lengthy &3 mm.; length of wing 6*8 mm.; width of head 
2 5 mm.; width of front at vertex *8 mm. 

Habitat: Lorha, Gold Coast; Dr. J. F. Corson, 18.4.1914; one 
female. 

This prettily marked species does not seem to be closely related 
to any West African form. 

Haematopota angustifronSy n. sp. 

Dark medium-sized species with a relatively narrow front; 
dorsum of thorax dark sepia-coloured, with two pale admedian 
stripes on the anterior half, light grey posterior border and pale 
scutellum; dorsal surface of abdomen dark with traces of paired 
admedian spots on the apical segments. Wings warm sepia- 
coloured with coarse creamy-yellow markings. Legs dark, tibiae 
with the usual pale bands, front tibiae slightly dilated, hind tibiae 
uniformly thickened. 

Head (fig. 6): Face , jowls and basioccipital region light grey 
clothed with glistening yellowish hairs; face with two circular black 
spots, one immediately below the base of each antenna. Front 
narrow, its length about one and five-eighths times the width at the 
vertex, dark velvety brown in colour with very narrow yellowish- 
brown lateral margins extending to encircle the frontal spots; vertex 
with a darker brown, more or less triangular, blotch divided in the 
middle line by a fine pale stripe; front clothed with short black 
hairs, sparse golden-brown hairs on the paler areas. Three frontal 
spots present, the median small but distinct, the laterals large, some¬ 
what rhomboidal in shape, and either narrowly separated from or 
in contact with the eyes. Frontal callus deep, shining, almost 
black, not extending from eye to eye, prominent; upper margin 
rounded but slightly depressed on each side of the middle line, 
lower margin concave above the antennae. Dark spot below callus, 



186 


quadrate, conspicuous. Antennae (fig. 6 j first joint dark yellow 
becoming yellowish-grey towards the apex, long and cylindrical, 
not incrassate, clothed with black hairs; second joint similarly 
coloured, its upper angle very slightly produced ; third joint 
elongate, dark brown to dark yellowish-brown, paler, yellowish- 
brown at extreme base; annuli distinct, almost black, the third very 



Fig. 6. Hatmatopota angustifrons , n.sp., tide view of head, 
female, (x 18, about). 

little shorter than the two preceding taken together. Palpi first 
segment greyish-yellow, clothed with long glistening cream-coloured 
hairs ventrally and laterally, and sparsely with long black hairs 
dorsally; second segment, except a large area on the outer side, 
greyish-yellow, clothed with long cream-coloured hairs, outer side 
mostly dark grey covered with short black hairs. 

Thorax: Admedian stripes on dorsum, creamy white, extending 
nearly to the inner extremities of the transverse suture, each followed 
immediately below suture by a conspicuous, triangular, similarly 
coloured, spot; crescentic marks on posterior margin, creamy-white, 
sharply defined and relatively large; dorsum clothed with golden- 
yellow and minute black hairs intermixed, those of the former colour 
more numerous on the sides of the anterior region and longer and 
denser on the posterior region; humeral calli mouse-grey clothed 
with long whitish hairs. Scutellum creamy-white with dark hind 





border, the pale area prolonged in the middle line in some specimens 
to the apex; clothed with creamy-white hairs. Pleurae greyish- 
brown above, slate-grey below, clothed with pale yellowish hairs. 

Abdomen: Slightly darker than the thorax with yellowish hind 
margins to the segments and a pair of ill-defined, elongated, dull 
yellowish, admedian spots on the fourth, fifth and sixth tergites; 
clothed with short black hairs on the main portion, yellowish hairs 
on the paler hind margins and rather long, dense, glistening white 
or yellowish-white hairs on the extreme lateral apical corners. 
Venter greyish-brown darkening towards the apex, with hind 
margins yellow but becoming greyish laterally; clothed with 
glistening yellowish hairs interspersed with black hairs in the central 
region of the penultimate segment, and with erect black hairs on the 
last segment. 

Wings (Plate XIII, fig. 2) : Pale markings coarse, creamy-yellow 
and conspicuous; rosettes large and distinct; apical sinuous mark 
relatively broad in its upper and lower portions but greatly 
narrowed and sometimes interrupted near the middle of the apical 
cell; posterior cells, except the fourth, With very large pale blotches 
along the wing margin, that in the first being formed by the lower 
extremity of the apical mark; axillary cell with a broad clearly- 
defined pale band running from the basal loop round the proximal 
margin and joining a somewhat sinuous pale line which extends 
upwards to the fifth longitudinal vein, and with a large pale blotch 
in its distal angle which extends well into the proximal angle of the 
fifth posterior cell. Stigma elongate, conspicuous, brownish-black, 
the extreme basal portion yellowish; veins brownish-black. Squamae 
brown. Halteres entirely pale cream-coloured. 

Legs: Front pair black, middle and hind pairs dark brown. 
Coxae of the first pair of legs dark brown or black with the proximal 
fourth greyish; clothed with long white hairs basally and with black 
hairs on the remaining portion. Femora, front pair clothed with 
black hairs which are rather long on the dorsal surface; middle and 
hind pairs clothed with black and occasional white hairs, those on 
the ventral surface of the hind pair being somewhat longer and more 
noticeable than usual. Front tibia with a broad white basal band, 
middle and hind tibiae each with two yellowish bands, the distal 
bands on the hind pair being generally incomplete, narrower and 



188 


darker; greater part of tibiae clothed with dark hairs, pale bands 
with white or yellowish hairs, interspersed on the hind pair with 
black hairs—dorsal surface of hind tibiae with rather long dense 
black hairs. Tarsi of middle and hind legs with the first segments 
narrowly cream-coloured at the base; clothed with black hairs. 

Length , 8‘8-io mm.; length of wing 8*5-9 nun.; width of head 
3*25-3*5 mm.; width of front at vertex ’65-75 mm. 

Habitat: Belgian Congo; Dr. E. Dutton and Dr. J. L. Todd, 
1904; six females. 

The exact position of H. angustifrons is somewhat difficult to 
define. In a general way it resembles H. grahami , Aust., and 
displays affinities to this species in the narrowness of the front, the 
shape of the frontal callus and, to a certain extent, in the arrange¬ 
ment of the pale markings on the body. The most striking character 
of H . grahami , however, namely the presence of fringes of long hair 
on the femora and hind tibiae, is not present, although the hairs on 
these parts in H . angustifrons are perhaps more conspicuous than 
usual. 


Haematopota exiguicornuta , n. sp. 

Small obscurely marked species with extremely short antennae; 
dorsal surface of thorax dull brown with three ill-defined, pale, 
longitudinal stripes; dorsum of abdomen slightly darker than that 
of the thorax, with inconspicuous admedian paired spots on some 
of the segments. Legs somewhat pale, the tibiae ringed. Wings 
with rather coarse pale markings and inconspicuous stigma. 

Head (fig. 7): Face y jowls and basioccipital region light grey, 
clothed with white hairs; upper part of face, at the side of each 
antenna, dark brown with a narrow black lower border and with 
yellowish pubescence. Front sepia-coloured, slightly paler round 
the lateral frontal spots and along the margins of the eyes; clothed 
with short dark brown hairs. All the frontal spots present but the 
median spot much reduced and scarcely visible under the pocket 
lens, lateral frontal spots separated from the eye on each side, 
moderately conspicuous, and more or less diamond-shaped, each 
with its longer axis pointing inwards and downwards. Frontal 
callus in the form of a narrow transverse dark reddish-brown band 
extending nearly from eye to eye; upper and lower margins gently 
curved and almost parallel, the former slightly concave. Spot 



189 

below callus small dark brown and divided in the middle line. 
Antennae (fig. 7) pale, shorter than the head; first joint short, 
dark reddish-brown, partly shining and swollen, clothed with black 
hairs; second joint paler brown, its upper angle considerably 
prdduced; third joint very short, yellowish, considerably flattened 
from side to side, the basal portion almost as broad as long; 
annuli (at least the first and second) indistinctly separated, the last 
not as long as the two preceding taken together and darker brown 


H.S.L. 

Fig. 7. Haematopota exiguicornuta , n.sp., side view of head, 
female, (x 18, about). 

at apex. Palpi mouse-grey; first segment clothed with long white 
hairs, except dorsally and apically, where black hairs occur; second 
segment somewhat dilated basally and gradually tapering to a 
bluntly rounded apex, clothed with long glistening white hairs on 
the ventral surface, and on the outer side with shorter black hairs. 

Thorax : Longitudinal greyish stripes on dorsal surface 
inconspicuous, the median stripe extending nearly to the scutellum, 
the admedian stripes reaching almost to the transverse suture, each 
with an indication of the usual spot below its posterior extremity; 
crescentic marks on posterior margin faintly marked, hardly 
recognisable; sides slightly paler; dorsum sparsely clothed with 
short glistening white and black hairs intermixed. Scutellum dull 
brown with greyish hii\d margin, clothed with hairs similar to those 
on the main portion of the dorsum. Pleurae grey, clothed with 
glistening whitish hairs. 

Abdomen : Hind and lateral margins of tergites narrowly light 
grey, the pale area being comparatively broad on the sides of the 
first and second segments; ill-defined admedian spots present on 
the second to seventh tergites inclusive; dorsum clothed mainly 




190 


with black hairs, pale margins with white hairs. Venter similar 
to dorsum, but without admedian spots, with the first segment grey 
and with the white hairs more widely and irregularly distributed. 

Wings : Pale sepia, the light markings somewhat indistinct and 
arranged as shown in Plate XIII, fig. 5. Stigma very faintly 
marked, the distal two-thirds pale brown, the proximal third 
whitish; veins reddish-brown. Squamae sepia-coloured. Halteres> 
stems cream-coloured, knobs dark brown with a relatively broad 
creamy stripe along the middle line. 

Legs : Pale brown. Coxae grey, clothed on the apex with 
black hairs and on the remaining portion with white hairs. Femora 
mostly greyish pollinose, clothed with black hairs, interspersed, 
especially ventrally and laterally, with white hairs. Tibiae pale 
brown, the front pair with a pale basal ring, the middle and hind 
pairs each with two pale rings; clothed chiefly with black 
hairs, sparse white hairs (which are more numerous on the hind 
tibiae) being present on the pale rings. Tarsi, front pair dark 
brown, middle pair yellowish-brown, hind pair rather paler than 
front pair, each with the base of the first segment yellowish; all 
clothed with black hairs. 

Lengthy 6 mm.; length of wing, 6*2 mm.; width of head, 
2 5 mm.; width of front at vertex, 0*7 mm. 

Habitat : Parade ground, Lokoja, N. Nigeria; Dr. W. M. 
Manuk, 19.7.1910; one female. 

The peculiar and characteristic antennae of H. exiguicornuta 
render its identification, and separation from species which resemble 
it in general facies, a comparatively simple matter. 

Haematopota corsoniy n. sp. 

Small dull sepia-coloured species with short antennae and 
relatively large dark areas on the face; dorsum of abdomen with 
indications of paired spots and with narrow pale hind margins to 
the segments. Wings grey, the pale markings somewhat reduced 
and inconspicuous; stigma well marked. Legs pale brown, tibiae 
unbanded. 

Head (fig. 8) : Face light grey with a conspicuous black area 
extending between the antenna and the eye on each side, narrowly 
separated at its extremities from both, and with the greater part 
of the central portion, below the antennae, dark-brown; jowls and 



basioccipital region light grey, clothed with long whitish hairs. 
Front , sepia with narrow yellowish-grey lateral margins extending 
partly round the lateral frontal spots. Median frontal spot absent 
or nearly so, lateral spots more or less quadrate and in contact with 
the eyes. Frontal callus narrow, shining, reddish-brown, almost 
divided by a longitudinal depression in the middle line and with 
the lateral extremities of its lower margin touching the eyes; upper 
margin nearly straight, curving downwards for a short distance in the 
centre, lower margin slightly concave above the base of each antenna. 
Spot below callus well-defined, dark brown, triangular, its apex 
directed downwards. Antennae (fig. 8) dark yellowish-brown, the 



Fig. 8. Haematopota corsoni , n.sp., side view of head, 
female, (x 18, about). 

tips (annuli) dark brown; first joint slightly incrassate and shining, 
clothed with short black hairs; second joint with the upper angle 
moderately produced; expanded portion of third joint short— 
slightly longer than broad—and somewhat flattened from side to 
side, terminal annuli taken together as long, or almost as long, as 
the expanded portion, last annulus equal in length to the first and 
second combined. Palpi grey, the outer side of the second segment 
with short black hairs. 

Thorax : Dorsum dull sepia, with three inconspicuous, smoke- 
grey, longitudinal stripes extending from the anterior margin to 
the region of the transverse suture, the spots below the posterior 
extremities of the admedian stripes very faintly marked; crescentic 
marks absent; sides, except a small area immediately behind each 
humeral callus, and scutellum dark; humeral calli and areas 
indicated grey, the former clothed with long white hairs; dorsal 
surface sparsely clothed with short whitish or yellowish-white 
hairs. Pleurae grey, clothed with white hairs. 



192 


Abdomen : Dorsum slightly darker than the jthorax, the tergites 
with very narrow brownish-grey hind margins, that on the second 
expanding into a small greyish triangle in the middle line; lateral 
margins of first to fifth segments greyish; paired, admedian spots 
on the second to sixth tergites, indistinct, brownish-grey in colour 
and approximated to the anterior margins. Venter generally rather 
darker than the dorsum, but with the whole of the first and second 
segments, except the central portion of the latter, mouse-grey; hind 
and lateral margins paler, the hind margins of the proximal scutes 
light grey. Both surfaces of abdomen clothed with white and black 
hairs, the white hairs limited to the pale margins of the tergites, 
but more widely and irregularly distributed on the stemites. 

Wings (Plate XIII, fig. 6): Pale markings white, inconspicuous, 
distributed as shown in the illustration. Stigma sepia, the extreme 
base pale. Squamae white, becoming yellowish towards the 
margins. Halteres , stems mostly cream-coloured, but darker at the 
base, knobs dark brown with a yellowish-brown central stripe. 

Legs : Coxae steel-grey, clothed with glistening white hairs. 
Femora, front and hind pairs dull pale-brown, middle pair some¬ 
what yellowish-brown; clothed with short black and white hairs 
intermingled. Tibiae similar in colour to the femora, without the 
usual pale rings but with a few short white hairs present on those 
positions where the pale bands generally occur, thereby giving the 
tibiae faint indications of bands. Tarsi, front pair slightly darker 
than the middle and hind pairs, the first segment of each of the 
middle and hind pairs faintly yellowish or yellowish-brown at the 
base. 

Lengthy 5*4 mm.; length of wing, 5 5 mm.; width of head, 
2 mm.; width of front at vertex, o*6 mm. 

Habitat : Salaga (15 miles North), Gold Coast; Dr. J. F. 
Corson (per Mrs. J. F. Corson), 17.10.1914; one female. 

This small and obscure species may, on a casual examination, 
easily be confused with Haematopota exiguicornutay n. sp. It may 
be readily distinguished, however, by the less swollen first antennal 
segment and the longer and more distinctly separated terminal 
annuli of the third joint, by the more conspicuous stigma and 
different arrangement of the pale markings on the wings, and by 
the unbanded tibiae, as well as by certain other, less obvious, 
differences. 



*93 

WEST AFRICAN SPECIES OF HABMATOPOTA 

Since three of the species of Haematopota described above were 
obtained in West Africa, a tabular statement of the more important 
distinctive characters of those members of the genus recorded from 
the British Possessions on this coast may prove of some value to 
medical officers and others interested in the biting flies of this 
region. The following table, therefore, refers to the females of those 
species at present known to occur in Gambia, Sierra Leone, Gold 
Coast or Nigeria, and its preparation has been greatly facilitated by 
Mr. E. E. Austen’s excellent descriptions of his numerous species. 

1. Front and hind femora and tibiae fringed with long coarse hair ... 2 

Front and hind femora and tibiae not fringed with long coarse 
hair. 3 

2. Hind tibiae greatly swollen with two ill-defined pale rings; frontal 

callus widely separated from the eyes; grey thoracic markings 
extensive . bullatifrons, Aust. 

Hind tibiae scarcely swollen with one broad pale ring; frontal 
callus almost touching the eyes; grey thoracic markings not 
extensive . grahami , Aust. 

3. Legs ornate with pale rings on the tibiae (distinct on, at least, the 

front pair). 4 

Legs inornate or with obscure rings on the tibiae. 12 

4. Hind tibiae with one pale ring . 5 

Hind tibiae with two pale rings . 6 

5. Blackish species with extensive grey thoracic markings, conspicuously 

banded tibiae and pronounced milky-white wing markings 

decora. Walk. 

Dusky species with dark wings, narrow' grey thoracic markings and 
less distinctly banded tibiae . hastata , Aust. 

6. Anterior tibiae with two pale rings (the distal ring often faint) ... 7 

Anterior tibiae with one pale ring . 8 

7. Larger (10-12 mm.) reddish-brown species; first antennal segment 

distinctly swollen; abdomen with a rather broad median stripe, and 
with trace* of spots on the terminal segments ... vitlata , Loew. 

Smaller (9*6 mm.) sepia-coloured species; first antennal segment not 
swollen; abdomen with a narrow median stripe and large spots on 
all segments. puntens, Aust. 

8 . Antennae longer than the head . 9 

Antennae very short and inconspicuous, shorter than the head (basal 
portion of third joint greatly expanded) ... exiguicornuta , n.sp. 




i 9 4 


9- First joint of antenna not distinctly swollen . io 

First joint of antenna distinctly swollen and shining, small dark species 
with conspicuously-marked wings. pinguicomis , n.sp. 

10. Brown species; background of wings dark, the light markings 

distinct .11 

Greyish species; background of wings very pale, the light markings 
indistinct . fallidipmnis , Aust. 

11. Scutellum brown; first joint of hind tarsi partly yellowish. 

torqums , Aust. 

Scutellum grey ; first joint of hind tarsi entirely brown. 

cordigera , Bigot. 

12. Brownish-yellow or greyish species with conspicuous stripes on the 

thorax .13 

Dark brown species without conspicuous stripes on the thorax (but cf. 
tcnuicrus, Aust.) .15 

13. Central part of wing hyaline or semi-hyaline, only the tips and hind 

border infuscated and with inconspicuous pale markings ... 14 

Wings sepia with clearly defined pale markings, extending to the base 

pertintns y Aust. 

14. Brownish-yellow species ; frontal callus yellowish ... beringcri, Aust. 

Greyish species; frontal callus dark brown ... semiclara , Aust. 

15. Anterior tibiae not dilated .16 

Anterior tibiae distinctly dilated; small dark species with first joint of 
antenna swollen . lacessens , Aust. 

16. Antennae moderately elongate ; basal portion of third joint slender 17 

Antennae very short, scarcely longer than the head; basal portion of 
third joint broad . corsoni , n.sp. 

17. Thorax conspicuously striped, abdomen indistinctly spotted; first 

antennal segment very short and somewhat swollen... tenuicrus, Aust. 

Thorax inconspicuously striped, abdomen unspotted; first antennal 
segment longer and not swollen. gracilis , Aust. 



*95 


REFERENCES 


Austen, E. E. (1909). Illustrations of African Blood-Sucldng Flics. 

- (1908 and 1911). New African Phlebotomic Diptcra in the British Museum 

(Natural History), Parts I, II, III, IV and VIII Tabanidae. Ann. Mag. Nat. Hist., 
8th Series, Vol. I, pp. 209 and 401; Vol. II, pp. 94 and 274; Vol. IX, p. 353. 

- (1912 and 1914). New African Tabanidae, Parts II, III, IV. Bull. Ent. Res., 

Vol. Ill, pp. 329 and 399; Vol. IV, p. 283. 

Loew, H. (i860). Dipt.-Fauna Sudafrika’s, Part I. 

Ricardo, Miss G. (1906). Notes on the genus Haematopota of the family Tabanidae in 
in the British Museum Collection. Ann. Mag. Nat. Ilist., 7th Series, Vol. XVIII, 
P* 94 - 

Speiser, P. (1910). Orthorhapha. Zool. Kilimandj.-Meru Exped., 1905-6. Diptcra, 
Band II, Abt. 10, p. 31. 

Surcoup, Jacques. (1908). Description de Diptires piqueurs Africains, Tabanides du genre 
Haematopota. Bull. Mus. Nation. d’Hist. Natur., Vol. XIV, p. 153. Description 
d’un Tabanidae (Haematopota) d’Abyssinie. Ibid., p. 224. 

- (1909). Note pr 61 iminaire sur la syst&natique du genre Chrysozona. Description 

de deux genres nouveaux. Ibid., Vol. XV, p. 453. Description de Tabanides 
nouveaux. Ibid., p. 537. 


Surcouf, J. M. R., and Ricardo, Miss G. (1909). Etude Monographique des Tabanides 
d’Afrique. 

Surcouf, J. M. R. (1911-1912). Note sur les Tabanides du Congo Beige des Mue6cs de 
Bruxelles et de Tervueren. Rev. Zool. Afric., Vol. I, p. 22. Deuxi£me note sur les 
diptires piqueurs du Mus6e du Congo Beige. Ibid., p. 86. 



196 


EXPLANATION OF PLATE XIII 

Fig. r. Wing of Haematopota pinguicornis, n. sp., female (x 7A). 
Fig. 2. Wing of Haematopota angustifrons, n. sp., female (x 7). 
Fig. 3. Haematopota transvaalertsis, n. sp., female (x 3). 

Fig. 4. Haematopota theobaldi, n. sp., female (x 3). 

Fig. 5. Wing of Haematopota exiguicormtta, n.sp., female ( x 7-J). 
Fig. 6. Wing of Haematopota corsoni, n. sp., female (x 7^). 

NOTE.— In figs. 1, 2, 5 and 6 the background of the wings has 
been darkened in order to render the light markings more 
conspicuous and to overcome halation. 



V* 


Annals Trop. Med. of Parasitol Fo/. /A' 


PLATE XIII 



i 


2 




C. Tinling Co., Ltd. Imp. 



'97 


EXPERIMENTS WITH SALVARSAN- 
COPPER IN TRYPANOSOMIASIS 

BY 

HARALD SEIDELIN, M.D. 

(Received for publication , 23 February , 1915) 

With Table 

Just before I left for West Africa, in July, 1913, I received from 
Geheimrat Ehrlich a supply of a new preparation, K 3 , salvarsan- 
copper, which he asked me to use in the treatment of yellow fever 
cases. Unfortunately I had no opportunity of complying with this 
request, because I saw only yellow fever cases of a mild type, in 
which an indifferent treatment was obviously all that was required. 
Several severe cases occurred during my stay in Lagos, but they 
were either convalescent when they came under my observation, or 
they died almost immediately after admission to hospital. 

It was also my intention to try the treatment in experimental 
yellow fever infection in guinea-pigs, but here again the conditions 
were unfavourable. The severity of the experimental infection 
varied considerably, and it was never possible to tell whether an 
animal would survive or succumb to the disease. Experiments of 
this nature would have to be conducted with large parallel series of 
infected and non-infected animals, and such experiments we were not 
in a position to undertake. 

Under these circumstances I resolved to test the value of K 3 in 
trypanosome-infections. For this purpose, Dr. J. W. Scott Macfie 
kindly supplied me with a strain of a trypanosome ( T . brucei group) 
which he had for some time kept in guinea-pigs and rats. 

The results of my experiments are given in the accompanying 
table, which requires but little explanation. The animals experi¬ 
mented upon were white rats; the infection was transmitted by means 
of intraperitoneal inject tons of a few drops of blood drawn from the 
cavernous sinus, according to the technique described by Pettit 
(1913). The trypanosome strain proved very virulent, the infected 
animals, when not treated, dying from eight to seventeen days after 
inoculation. 

Geheimrat Ehrlich had advised me to start with doses of 0 05 or 



198 


o io gramme in experiments upon human patients. The average 
weight of the rats was about 60 grammes, and the variations in 
weight were small; thus, the proportion in weight of rat to man 
would be approximately 1 : 1,000. Accordingly, in the first 
experiment I took 0*0001 as a medium dose, using in addition two 
higher and two lower doses. As no satisfactory result was obtained 
in this experiment I increased the dosage, always proceeding by 
multiples of 2. It will be seen from the table that no striking result 
was obtained until I reached a dose of eight times the one recom¬ 
mended, and that the best results, as far as the prolonged absence 
of the parasites from the blood was concerned, were obtained by the 
injection of 0*0064 gramme instead of 0 0001 gramme. In such a 
case the trypanosomes, which had been very numerous before the 
injection, were absent from the blood on the following day and 
remained absent for fifteen days; the animal died twenty-eight days 
after the inoculation. In several other cases I succeeded in 
prolonging the life of the animals for a few days more, although the 
parasite-free interval was of shorter duration. It appears probable 
that repeated injections of smaller quantities would have given better 
results, but this method of treatment was not fully investigated. The 
number of animals available was not sufficient for work on different 
lines, and I therefore confined myself to the investigation, in the 
first instance, of what I regarded as the problem of primary 
importance, namely, the possibility of establishing a real ‘ therapia 
sterilisans magna' In this I failed : a single injection, however 
large, did not kill all the parasites in an infected animal, though it 
made them disappear, for a considerable length of time, from the 
peripheral blood. It would hardly be possible, even in experiments 
on animals, to use larger doses- than those I employed, as several 
animals died a comparatively short time after the injection without 
reappearance of the parasites, and in others a more or less extensive 
necrosis of the tissues was produced at the site of injection. 

The results may be discussed in a few words. 

In series A very little effect was obtained. No appreciable 
numerical reduction of the parasites was observed, even after the 
injection of 00004 gramme K 3 , but in all cases the parasites 
appeared much less lively on the day following the injection than 
they had been the day before and as compared with the parasites in 



199 


the blood of the control animals on the same day. The longest 
period of survival obtained in the two animals which had received 
the largest doses of K a . 

In series B the action of the drug was evident. Two animals 
survived the inoculation for 28 and 29 days, respectively, and in one 
of them trypanosomes were absent from the blood for six days; this 
animal had received an injection of 0 0016 gramme K 3 . 

In the two previous series only the doses administered had been 
varied, but in series C the drug was injected into animals which 
were in different stages of infection. In three animals, as in all 
those belonging to the series A and B, trypanosomes were numerous 
at the time of injection, but in the other three they had not yet 
appeared in the peripheral blood; one animal of the latter group had 
been inoculated about 75 minutes before the therapeutical 
injection. The stage of infection proved to be of little importance. 
The hope had been entertained that a dose, which showed a marked 
effect when numerous trypanosomes were present in the blood, might 
entirely destroy an infection in its initial stage; but this hope was 
disappointed. In rat 232, trypanosomes did not appear in the 
peripheral blood until eight days after inoculation; this free interval 
may be assumed to be composed of two periods, namely, one 
corresponding to the action of K 3 , followed by another corresponding 
to the stage of incubation. The dose administered was 0 0016 
gramme. The same dose was given in the cases 199, 220, and 229, 
in which cases the parasite-free intervals varied from four to six 
days. Thus, if we reckon five days for the first period in rat 232, 
we get three days for the second period, which is precisely the usual 
duration of the incubation stage. In the case of rat 221, trypano¬ 
somes reappeared in the peripheral blood on the tenth day after the 
injection of 0 0032 gramme K 3 ; on the same day 0 0064 gramme was 
injected, whereupon the trypanosomes again disappeared from the 
blood and remained absent until the animal died, twenty-three days 
after inoculation. 

In series D the doses were increased. The parasite-free intervals 
were prolonged, the maximum of fifteen days after a single 
injection being attained in the case of rat 241, which received 0 0064 
gramme K 3 ; but the animals did not survive as long as those in 
experiment C. 



200 


In series E repeated injections were given, the dose in each 
instance being 0 0064 gramme; in this way a parasite-free period of 
twenty days was obtained, but the length of survival was not 
increased. 

In several of the experiments of the series D and E the animals 
died without the trypanosomes having reappeared in the blood. 

The experiments in series F were undertaken in order to test the 
possibility of producing a K 3 -resistant strain of the trypanosome. 
Two rats were inoculated with blood from an animal in which the 
parasites had first disappeared after the injection of K 3 , and after¬ 
wards reappeared. One of the animals was injected twice with K 3 , 
and the effect of the drug was considerably less than in the other 
experiments. 

The results here reported are interesting from a general point of 
view, and leave no possible doubt as to the powerful action of 
salvarsan-copper upon this strain of trypanosome, but they are not 
conclusive with regard to the therapeutical value of the drug. After 
having concluded my experiments, I became acquainted with the 
paper by Van den Branden (1913) on the effect of K 3 in human 
trypanosomiasis. His results are far superior to those obtained by 
me in rats and with a different species of Trypanosoma \ this confirms 
the information given me by Professor Ehrlich that K 3 has a 
markedly different effect upon different trypanosomes. 

The experiments here recorded were carried out whilst I was 
working for the Yellow Fever (West Africa) Commission of the 
Colonial Office, at the Medical Research Institute, at Yaba, near 
Lagos, Nigeria. To the Director of the Institute, Dr. A. Connal, 
I am indebted for the facilities afforded me. 


REFERENCES 


Branden, F. van den (1913). Note prelimimire sur quelques cjsais de traitement dc la 
Trypanose Humaine par Salvarsankupfer. Arcb. f. Scb. u. Tropen-Hyg ., Leipzig, 
Dec., XVII, 24, pp. 845-849. 

Pettit, A. (*913)* Proc&te simple pour pr£lever du sang cher les pstit* rongeurs. 
C. R . Soc. Biol., Paris, 10 jan., LXXIV, 1, pp. 11-12. 



Result 


Date 

inoct 


[0. 1 

tio 

1 

1 9 *- 

1 

1611 

2: .S 


2Z> 

1 

163 | 

22.? 

1 

164 | 

| 21.? 

169 

I 22.? 

165 

| 22.) 

l66 

1^ 

167 

22.1 

(68 

22.1 

*59 

, 1 ill 

i6e 

, Hi 




*7 


18 


*9 


Death 13 days after inoculation 

i> 7 n 

n *4 m 

*i 8 

» 11 » 

u 13 » 

99 14 n 

11 *7 9* 

n *7 »> 

» 7 n 

n 14 99 


Spleen very much enlarged 
Peritoneal infection 

Spleen very much enlarged 



220 | 

28. 

221 

1 28, 


229 L 

230 Z 

23» 3 

2 3 2 4 


Death 12 days after inoculation j 


yy 

*3 


yy 

n 

*3 

yy 

yy 


29 

yy 

yy 


28 

** 

yy 

yy 

*5 


yy 


Death 28 days after inoculation 

?? -3 

M 22 

yy 3 2 1? yy 

>5 33 v V 

91 22 „ 

[p.T.O. 















200 


In series E repeated injections were given, the dose in each 
instance being 0 0064 gramme; in this way a parasite-free period of 
twenty days was obtained, but the length of survival was not 
increased. 

In several of the experiments of the series D and E the animals 
died without the trypanosomes having reappeared in the blood. 

The experiments in series F were undertaken in order to test the 
possibility of producing a K 3 -resistant strain of the trypanosome. 
Two rats were inoculated with blood from an animal in which the 
parasites had first disappeared after the injection of K 3 , and after- 
wards reappeared. One of the animals was injected twice with K 3 , 
and the effect of the drug was considerably less than in the other 
experiments. 

The results here reported are interesting from a general point of 
view, and leave no possible doubt as to the powerful action of 
salvarsan-copper upon this strain of trypanosome, but they are not 
conclusive with regard to the therapeutical value of the drug. After 
having concluded my experiments, I became acquainted with the 
paper by Van den Branden (1913) on the effect of K 3 in human 
trypanosomiasis^ His results are far superior to those obtained by 
me in rats and with a different species of Trypanosoma ; this confirms 
the information given me by Professor Ehrlich that K 3 has a 
markedly different effect upon different trypanosomes. 

The experiments here recorded were carried out whilst I was 
working for the Yellow Fever (West Africa) Commission of the 
Colonial Office, at the Medical Research Institute, at Yaba, near 
Lagos, Nigeria. To the Director of the Institute, Dr. A. Connal, 
I am indebted for the facilities afforded me. 


REFERENCES 

Branden, F. van den (1913). Note preliminaire aur quelques e*sais de traitement de U 
Trypanose Humaine par Salvarsankupfer. Arch . /. Scb . u. Tropen-Hyg ., Leipzig, 

Dec., XVII, 14, pp. 845-849. 

Pettit, A. (R913)* Proc6d6 simple pour pr£lever du sang che* les petit* rongeurs. 
C. R . Soc . Biol ., Paris, 10 jan., LXXIV, 1, pp. 11-12. 





Result 


Date t 
Rat inoculi 
No. tion. 



7 'J 

l6 

*7 

1 

18 

*9 

20 



161 

22.X 

... j 



! 

f 

... , 

Death 13 days 

. 1 

after inoculation 1 

Spleen very much enlarged 

162 

22. X 

..... 1 1 

... , 

1 1 


n 

7 

» 

- 

Peritoneal infection 

* 6 3 

22.X * 




1 

- | „ 

*4 

ff 

n 


164 

22.X - 




i ff 

1 

8 

ff 

tt 

Spleen very much enlarged 

169 

22.X 




1 

ff 

11 

ff 

tt 

tt » 

165 

22.X - 





ft 

*3 

ff 

tt 

tt tt 

166 

22.X - 



... 


ff 

H 

ff 

tt 

tt tt ; 

167 

22.X 

... 




ff 

*7 

ft 

tt 

tt tt 

168 

22.X 





ff 

*7 

tt 

tt 

tt tt 

1 59 

22.X 





ff 

7 

tt 

tt 

a a 

160 

22.X - 





ff 

*4 

tt 

tt 

tt tt 









1 95 

.5x1 - 

... 

... 

... 

1 

Death 

12 days after inoculation 


196 

15-Xl 






*3 

tt 

tt 


*97 

15.XI 






13 

tt 

tt 


198 

1 1 5.XI + + + 

-f 4 ' 4 * 




ff 

29 

tf 

tt 



) 1S-XI+ + + 

+ 4 -+ 




ff 

28 

tt 

tt 


t 9 J 

1 ' 5 X 1 





\ ff 

*5 

ff 

tt 



220 

28.XI ••• 


... 



Death 28 days after inoculation 

221 

28 .xi - 


... 



a 23 

229 

I.X1 - 

... 


... 

: l 


tt 22 

23O 

2.X! - 

1 

... | ... 

1 

n 32 tt a 

23 * 

. 3 -xi - j - 

j 


tt 33 

232 

4 * - ; - 



» 22 „ 


[p.T.O 














240 

8.XII 

1 

13.XII 



... 

... i 

Death 11 days after inoculation 

I 

2 4* 

8.XII 

13.XII 





2 ^ 

Necrosis at the site of injection 

2 43 

10.XII 

... 





»» *7 11 »» 

Large necrotic cavity at the site 
of injection 

2 44 

10.XII 

13XH 

* 



... 

ji 3 11 


2 47 

n.XII 

13.XII 




... 

4 n »> 


2 5° 

12.XII 

13XII 

1 




1 

1 _ 

10 ?» 

Spleen very slightly enlarged 


3o;XII 

4.I.14 


... 

... 


Death 19 days after inoculation 

3a.XlI 

4 I «4 


1 

1 

inj. of 
0-0064 g. 

n 2 ^ » »» 

# 


?appearcd ; subsequent treatment with K 8 .) 


22 7 

i.XII 

4.XII 

- 4 - 

+ + + 

+ + + 

4 - 4 - + 

Death 25 days after inoculation 

228 

1.XII 





... 

*3 )» »> 






201 


STUDIES IN BLACKWATER FEVER 

III.—THE RELATIONSHIP OF QUININE 
TO BLACKWATER FEVER* 

BY 

J. W. W. STEPHENS 

SIR ALFRED JONES PROFESSOR OF TROPICAL MEDICINE, UNIVERSITY OF LIVERPOOL 

AND 

W. STOTT 

HON. STATISTICIAN iO THE LIVERPOOL SCHOOL OF TROPICAL MEDICINE 

(Received for publication 24 February , 1915) 

In the literature of blackwater fever contradictory statements are 
made concerning the role of quinine, some observers asserting that 
quinine induces the attack of blackwater, others denying this. The 
solution of the question is not advanced by the affirmation or denial 
of supposed authorities. What we have tried to find out is whether 
a statistical examination of the times at which quinine is given 
and blackwater occurs respectively, reveals any relationship between 
the two. 

In examining the literature, an initial difficulty confronted us, 
viz., the incompleteness of the records. For our purpose it was, 
as will be seen, necessary to know as exactly as possible at what 
hour quinine was administered and at what hour the black¬ 
water ensued. It was often impossible to ascertain these facts, 
and we would state clearly that the only data we have used are those 
in which the exact times of the giving of quinine and the occurrence 
of the rigor or blackwater are stated. 

We have thus not considered cases where the statements are not 
more definite than, e.g., ‘quinine was given the day before,* ‘the 
day of,* ‘shortly before the blackwater,* etc., etc., and also, of 
course, those cases could not be considered where it was stated that 
no quinine had been taken. 

As it was practically impossible to get records of cases with 

• Part I. Annals of Trop. Med . and Parasit. (1913). Vol. VII, p. 479, 

Part II. Ibid. (1914). Vol. VIII, p. 639. 




202 


a complete quinine history we were not able to study such, 
consequently we proceeded to examine whether any relationship 
exists between the time of the blackwater and the time of the 
last dose of quinine. 

In some cases the time of the rigor is stated, in others the time 
of the blackwater. Very probably the time of the onset of the rigor 
marks the real onset of the attack and is perhaps more accurate in 
that respect than the time of the passage of blackwater, for we have 
no accurate knowledge as to how long haemoglobinous urine can be 
retained, although it seems to be generally assumed that haemo¬ 
globinous urine is irritating to the bladder. 

The data then that concern us are (i) the time when the last dose 
of quinine was given; (2) the time of onset of the rigor; (3) the time 
of first passage of blackwater. 

We have obtained these or some of these data in 372* cases, the 
records of which we have examined. 

Time of quinine: This was given in 157 out of 372 cases. 

Time of rigor: This was given in 121 out of 372 cases. 

Time of blackwater: This was given in 217 out of 372 cases. 

It will be noted that it is possible either (1) that the 157 cases in 
which the time of quinine is given are none of them the same cases as 
the 121 cases in which the time of the rigor is given, or (2) that the 
157 cases (except 2) of quinine are none of them the same as the 217 
cases in which the time of the blackwater is given, but although both 
these possibilities could not hold good for all the cases in categories 
(1) and (2), they might do so for a part of both, so that in the 
first table the fields of observation are not (entirely) coincident. At 
present we disregard this. 

The data were now arranged as in Table I. We have taken 
eight three-hour periods, because it was (frequently) impossible to 
define the time more accurately, and also because if the day were 
divided up into twenty-four hours the number of cases falling under 
each hour would be too small whereon to base any deductions. 

On examining the columns of deviation from an average, it will 
be noticed that each has a very decided maximum. 

(1) Time of taking quinine. Upon the assumption that a random 

_ \ _ 

• Four cases have been excluded in which the interval between the quinine and the onset of 
the rigor or blackwater was greater than 24 hours. 



203 


distribution* is a priori to be expected, it can be calculated that the 
large deviation, 28 in excess of the average, would only be expected 
to occur once in over 10,000 cases, so we may fairly say 
that the distribution is not a random one, and that there is some 
reason for taking quinine between the hours of 6 a.m. and 9 a.m. 

This we can assume is due to our methods of life, the time of 
rising in the morning determining the time of taking quinine. 


Table I 


Hour 

Number 
of people 
taking 
quinine at 
stated hour 

Deviations 

from 

average of 
20 per 3 
hours 

Number of 
rigors 

occurring at 
stated hour 

Deviations 

from 

average of 
15 P cr 3 
hours 

Number of 
cases of 
blackwater 
at 

stated hour 

Deviations 

from 

average of 

28 per 3 
hours 


1 

12— 3 

3 

-17 

8 

- 7 

,8 

—10 


3-6 

18 

— 2 

7 

- 8 

15 

-«3 

a.m. J 




! 





6— 9 

48 

+ 28 

14 

— 1 

20 

- 8 


,12— 6 

26 

i + 6 

38 

+ 23 

47 

+ 19 


> 2—3 

12 

- 8 

1 

22 | 

+ 7 

5 ° 

+ 22 


3-6 

22 

+ 2 

10 

~ 5 

37 

+ 9 

P.M. - 









6-9 

16 

! -♦ 

9 

- 6 

17 

-11 


1 9 — 12 

12 

- 8 

>3 

— 2 

1 

x 3 

-■5 


.* j 

121 

1 

2I 7 

1 


(2) The onset of rigor . The maximum deviation in this case is 
between the hours of 9 a.m. and 12 noon, Here we have 23 above 
the average, and it is also at least 10,000 to one that this does 
not occur at random, that is, there is some cause at work which 
determines the time. 

(3) Passing of blackwater . Here we have the maximum, 22, 


• By a random distribution in this case, we mean the distribution that would be expected to 
occur, if each case was entered at random in any of the eight divisions into which the day was 
divided, by a person who had no bias in favour of one hour rather than another, or if the division 
into which each case was entered was determined by picking a ball out of a bag containing eight 
differently coloured balls. Out of 157 cases we should expect about 20 in each of the eight 
divisions, but almost certainly there would be slight departures, owing to our random method of 
filling the divisions, but we have in our 6-9 a.m. division as many as 48 that is a departure from 
the average, 20, of 140 per cent. 




204 


between 12 and 3 p.m., but there is also a large positive deviation, 
19, between 9 a.m. and 12 noon. This would be expected to happen 
if the period between quinine and blackwater were a little over 
3 hours, many cases falling on the border line at noon. Again such 
a large deviation as 22 would only occur by chance once in 10,000 
cases. 

If now we move the figures for the rigor 3 hours back, the 
reason for so doing we shall presently see, and compare them with 
those of the quinine, we find that the coefficient of correlation ( r ) 
between them is r — 92 + *04, a very strong correlation (r = 1 
is perfect correlation). In non-mathematical language, correlation, 
positive or negative, means 1 fit/ and we may illustrate firstly the 
meaning of positive correlation in the following way: — 

Take sheets of paper in pairs, subject each pair to some operation, 
e.g., cutting with a pair of scissors through both at the same time; 
if we cut all by a straight line, it would be impossible, when the 
sheets were scattered, to bring the pairs together again, because 
many pairs would fit one another, but if we made a random 
zigzag cut through each pair, then it would be possible after 
separation to bring the pairs together, and the certainty we should 
feel that we had obtained a pair cut at the same time, would be so 
much the greater, the greater the number of zigzags and the greater 
their distinctive peculiarity (fig. 1). 

To illustrate secondly the meaning of negative correlation, we 
may similarly imagine a number of single sheets of paper cut across 
by a zigzag cut. When a fit is obtained between two pieces 
(belonging to an original whole) it is now a negative one, i.e., the 
peaks and depressions of one piece fit the depressions and peaks of 
the other. And here again the greater the number of peaks 
and depressions that fitted the greater the certainty that they 
belonged to an original whole (fig. 1). 

Now the method used in calculating the correlation coefficient 
gives due weight, both to the number of zigzags, and to the 
magnitude of the peaks or depressions. The symbol used for the 
correlation coefficient is r, and its numerical value varies from 
— 1 to + 1. r = o means no fit, r = + 1 means perfect positive 
fit, 7 = - 1 means perfect negative fit. 

Except in artificially chosen examples, we cannot expect perfect 



I. To illustrate correlation or fit.—There is a negative fit between the two pieces of 
paper A and A.i.—i.e., when A is inverted, as in the figure, the depressions of A fit 
the peaks of A.i and the peaks of A fit the depressions of A.i. There is a positive 
fit between the two pieces of paper I and 2—i.e., the peaks of I fit the peaks of 2, 
and the depressions of I fit the depressions of 2 . 




206 


fit, because we always have errors of observation, and errors due to 
random sampling, but the nearer we get to i, positive or negative, 
the greater our belief in a connection. 

Now the fit in the figures for quinine and rigor is very close, 
.92 + ’04 (instead of 1). As we said before, in obtaining a positive 
fit between two pieces of notepaper we should feel the more certain 
that they were cut at the same time the greater were the number of 
zigzags (and the greater their distinctive peculiarity). So that if 24 
zigzags fitted, the certainty would be greater than if there were only 
12 zigzags. Our cases, however, are insufficient to allow of our 
distributing the quinine and rigor figures over every hour of the 
day, i.e., into 24 observations, which would be desirable. 

We have said above that the fields of observation in Table I 
were not coincident, we now proceed to consider those cases only 
in which the fields are coincident, i.e., in each case the conditions 
affecting the taking of the quinine and the onset of blackwater are, 
so far as we can tell, identical. 

(1) We have 88 cases in which both the time of quinine and the 
time of rigor are given. 

Tabli II 




Number of 






people taking 

Deviations from 

Number of rigors 

Deviations from 

Hour 

quinine at 
stated hour 

average of 

11 per 3 hours 

occurring at 
stated hour 

average of 

11 per 3 hours 

| 

12— 3 

2 

- 9 

7 

“ 4 


3— 6 

IO 

: 

! 3 

- 8 

A.M. ^ 

| 

6— g 

3 ° 

+ 19 

9 

- 2 


' 9—12 

1 21 

+ IO 

! 

33 

+22 


t 12 3 

2 

- 9 1 

15 

+ 4 


3— 6 

11 

0 

6 

- 5 

P.M. - 

®— 9 

6 

: “ 5 1 

7 

~ 4 


' 9—12 

i 

6 

1 " 5 ! 

8 

- 3 

1 88 

i 1 

’ 88 

i 



The correlation between these deviations as they stand is r = 34, 
but when the figures for the rigor are moved three hours back the 
correlation is r = *85. 





207 



IZ -3 3-6 6- 9 9-/2 JZ' 3 3-6 6-9 9 ~/z 


F Lee so n. del 

Chart I. Showing the relationship of quinine and rigor in 88 cases. The mode 
of quinine is at 6—9 a.m., that of the rigors is at 9 a.m.—12 noon. 











208 


(2) We have 103 cases in which both the time of quinine and the 
time of blackwater are given. 


Table III 


Hour 

Number of 
people taking 
quinine at 
stated hour. 

Deviations from 
average of 

13 per 3 hours 

Number of cases j 
of blackwater 
occurring at 
stated hour 

Deviations from 
average of 

13 per 3 hours 


1 l2_ ~ 3 

1 

—12 

4 

“ 9 

A.M. -j 

; 3-6 

12 

— 1 

3 

—10 

1 





j 

6- 9 

35 

+22 

9 

- 4 

1 

L 9 — 12 

26 

+ 13 

30 

+ 17 

1 

l ^ 3 

4 

- 7 

21 

+ 8 

1 

1 3- 6 

*3 

0 

22 

+ 9 

P.M. J. 







J 6 — 9 

8 

- 5 

7 

- 6 


l 9—12 

4 

“ 9 

7 

- 6 



103 


i °3 



The coefficient of correlation of the deviations as they stand is 
7 = ‘34, but when the figures for the blackwater are shifted three 
hours back the correlation is increased to r = '95 (perfect correla¬ 
tion, 7 = 1). 

The increase in the correlations from 34 to ‘85 in one case and 
from *34 to ‘95 in the other, when the figures for the effect (rigor or 
blackwater) are moved through three hours to fit the (presumed) 
cause (the quinine) is very significant, and with such a high correla¬ 
tion, ‘85 in one case and 95 in the other, it is highly probable that 
there is a connection between the quinine and rigor or the quinine 
and blackwater, but it is still possible that there is none. 

For it must not be assumed that perfect correlation means that 
we have obtained a p 7 oof that the events are connected as cause and 
effect. 

(1) It is possible, for all we know to the contrary, that the mode* 

# By a mode is meant the most frequent occurrence of any particular fact or event—e.g., if 
the ages of six persons are 5, 5, 5, 5, 5 and 35 respectively, the average age is 10, but the 
mode is 5. 



209 



/ 2-3 3-6 6-9 9 - /2 / 2-3 3-6 6-9 9-/2 


P-Le fSON.^cl. 

Chart II. Showing the relationship of quinine and blackwater in 103 cases. The 
mode of quinine is at 6—9 a.m., that of blackwater is at 9 a.m.—12 noon. 













210 


(i.e., the most usual time) of the rigors in blackwater, from the 
nature of the disease, t occurs at 9 a.m.-i2 noon. If this be so, then 
of course a strong correlation might exist between it and any other 
(antecedent) phenomenon that had a mode at some antecedent time. 
For example, if we investigated the time of taking coffee we 
should probably find that the mode (i.e., the most usual time) was 
between 6 a.m. and 9 a.m., and so by moving the figures for the 
blackwater three hours back we should get a very strong correlation. 
It is necessary, then, to alter the conditions, either by altering the 
time at which quinine is taken or by observing a sufficient number 
of cases when quinine was taken in the afternoon, i.e., in this 
particular instance where there is no mode for coffee. If, then, we 
find that any alteration in the time of taking of quinine is followed 
by a similar alteration in the time of the rigor, the proof of 
connection is correspondingly strengthened. 

(2) It is possible that the modes which we have found do exist 
for quinine and rigor or blackwater, although they exhibit a high 
correlation, yet are not really related to one another as cause and 
effect. Both modes might be due to some other cause or causes; for 
instance, the rise in the price of wheat and the rise in the male death 
rate (between which there might be a strong correlation) might be 
due to the war, and yet we could not argue that the rise in the price 
of wheat was the cause of the rise in the male death rate. 

To sum up, we have really started with the assumption that the 
taking of quinine by persons with a certain diathesis is the cause, 
or a cause, of ‘blackwater.’ On this assumption we should expect 
to find a positive correlation between the time of taking the quinine 
and the symptoms of the illness. 

We do find such a positive correlation, and as the effect follows 
the cause usually after a certain period of time, we should expect the 
correlation to be brought nearer to unity as the figures for the effect 
are brought more in coincidence with the cause, in technical terms 

t We find, for instance, the following statements with regard to malaria in Mannaberg (1905):— 
‘ Maillot and the majority of observers since him state most paroxysms (about two-thirds of all 
cases occur, between midnight and midday—in other words, in the morning/ Moreover, according 
to Maillot, 4 the greatest number of quotidian parasites occur about 10 a.m. ; the smallest number 
between 9 p.m. and midnight/ Again, it is stated that * Maurel observed in Guiana the majority 
of paroxysms between 2 and 5 p.m/ Mannaberg himself states 4 that out of 107 cases 91 per cent, 
of the paroxysms occur during the period between 10 a.m. and 3 p.m/ We have not so far been 
able to examine these statements critically, so can form no estimate as to what they are worth. 



211 


as we bring the two modes together. This also we find. Had the 
quinine prevented blackwater we should have expected a large 
negative coefficient, and if the quinine had no effect we should have 
expected the correlation to be practically nothing. 

From what we have said, there are then three possible explana¬ 
tions of this correlation which we have found to exist between quinine 
and blackwater. 

(1) It is purely accidental, due to the fact that blackwater has a 
‘natural* mode in the 9 a.m.-i2 noon interval. This, as we have 
said, could be eliminated by seeing whether the same mode existed 
when quinine was taken at other times than 6-9 a.m. For this 
purpose we require further cases with the times accurately recorded. 

(2) The correlation is produced by some unknown cause or causes 
which determine both the fact that the mode of quinine is at 6-9 a.m., 
and that the mode of the rigors is at 9 a.m.-12 noon. We cannot 
suggest any such cause. 

(3) That the correlation is one of cause and effect. 

We believe, then, that, provided (2) can be excluded, that an 
examination of further cases, in the way we have done here, will 
decide whether (1) or (3) is the true answer to the question. 

By applying the statistical method adopted in this paper, we 
thought it possible that a solution of the question might be 
forthcoming, and although the cases we have been able to obtain are 
neither sufficiently numerous, nor recorded with the necessary 
accuracy of detail for us to claim that a solution has been arrived at, 
yet we believe that as a result of our analysis the problem can now 
be approached, when further cases are forthcoming, in such a way 
that we may fairly anticipate a solution in the near future. 

That this is a matter of vital importance, and not simply one of 
scientific interest, everyone who has any knowledge of the disease 
must realize. 



REFERENCES 


Broden, A. (1906). L’hemoglobinuric an Congo. Trav. Lab. Med. Leopoldville, Bruxelles, 
II, pp. 1-70. 

Campenhout and Dryepondt (1901). Fiivre bilie use hemoglobin urique. Trav. Lab. Med. 
Leopoldville, Bruxelles, I (1899-1900), pp. 51-117. 

Christophers, S. R., and Bentley (1918). Blackwater Fever. Scientific Memoirs by Off. 
Med. San. Dept. Govt. India, Simla, No. 35. 

Decks, W. E., and James, W. M. (1911). A report on hemoglobinuric fever in the Canal Zone. 
A study of its etiology and treatment. Pp. 177. With charts and tables. Mount 
Hope, C.Z. 

Doering (1898). Ein beitrag cur kenntniss der Kamerun-malaria u.s.w. Arb. a. d. k. Gesund- 
heitsamte, Berlin, XIV, S. 121 -137. 

Kleine, F. K. (1901). Ueber Schwarxwasserfieber. Zcitschr. Hyg. Inf., Leipzig, XXXVIII, pp. 
472-486. With charts. 

Koch, R. (1899). Ueber Schwarzwasserfieber (Ilamoglobinurie). Zeitschr. Hyg. Inf., Leipzig, 
XXX, pp. 295-327. 

Maillot (1836). .' Traite des fievres intermittentcs. Paris, 1836. 

Mannabkrg, J. (1905). Malarial Diseases. Nothnagel’s Encvcl. of Practical Medicine. 
English Edition, W. B. Saunders & Co., Philadelphia and London, p. 226. 

Maurel (1883). Traite des maladies paludiennes. Paris, 1883. 

Pampoukis, P. S., and Chomatianos, S. (1889). Recherches cliniques et experimentales sur 
Ph&noglobinurie quinique. Progria medical, Paris, VIII, 2 seric, pp. 3-6. 

Panse, O. (1902). Schwarzwasserfieber. Zeitschr. Hyg. Inf., Leipzig, XLII, 1, pp. 1-44. 
With charts. 

Plehn, A. (1896). Beitrage zur Kenntniss von Verlauf und Behandlung der tropischen malaria 
in Kamerun, Berlin. August Hirschwald. 

Plehn, A. (1914). Ein beitrag zur kenntniss der akut hamolytischen malaria (Schwarzwasscr 
fieber). Deutsch. Med. Woch., XL, pp. 1414-1416. 

Plehn, F. (1898). Die Kamerun-Kiiste. Studien zur Klimatologie, Physiologic und Pathologic 
in den Tropcn. Mit 47 Abbildungen und 1 Karte. Berlin, A. Hirschwald. 

Reports (1912). Reports on Blackwater Fever in the Tropical African Dependencies. 

Reports (1914). Blackwater in the Tropical African Dependencies. Reports for 1912. 
Schellong, O. (189 a). Die Malaria Krankheiten u.s.w. Berlin, 1890. 

Stephens, J. W. W., and Christophers, S. R. Reports to Malaria Commission of Royal Society, 
London. (1900) 1st Ser., pp. 12-42, pp. 42-75 ; (1901) 5th Ser., pp. 12-27; (1903) 
8th Ser., pp. 3-26. 

Tomaselli, S. (1897). La Intossicazionc clinica e. l’infezione malarica, Catania, pp. 
XVI +- 204. 



Volume IX 


June, 1915 


No. 2 


ANNALS 

OF 

TROPICAL MEDICINE AND 
PARASITOLOGY 


ISSUED BY 

THE LIVERPOOL SCHOOL OF TROPICAL MEDICINE 


Edited by 

Professor J. VV. W. STEPHENS, M.D. Cantab., D.P.H. 
Professor R. NEWSTEAD, M.Sc., J.P., F.R.S., A.L.S., F.E.S., Hon. F.R.H.S. 
Professor WARRINGTON YORKE, M.D. 

AND 

Professor Sir RONALD ROSS, K.C.B., F.R.S., M.D., F.R.C.S., 

Major I.M.S. (Ret.) 

Editorial Secretary 
Dr. H. B. FANTHAM, 

School of Tropical Medicine , 

The University , 

Liverpool . 



C. Tinting & Co., Ltd. 

Printers to the University Press of Liverpool 
53 Victoria Street 



21 $ 


GANGOSA IN NEW GUINEA AND ITS 

ETIOLOGY 


BY 

ANTON BREINL 

FROM THE AUITRAL1AN INSTITUTE OF TROPICAL MEDICINE 

(Received for publication 20 May , 1914) 

Plates XIV-XVII 

Gangosa, or Rhinopharyngitis mutilans, is a disease of limited 
geographical distribution, being, according to our present know¬ 
ledge, confined mainly to the Pacific Islands. 

According to Castellani and Chalmers (1910), cases of this 
disease were described first in 1828 by a Spanish Commission to the 
Marianne or Ladrone Islands, under the name of Gangosa, 
meaning ‘Nasal Voice.* Fordyce and Arnold (1906), and in the 
same year Leys, gave a clinical account of a similar disease in Guam, 
and were followed by Mink and McLean (1906) who discovered cases 
of Gangosa in the Ladrone and Caroline Islands. 

, . Later, Stitt (1907) reported the occurrence of Gangosa in a white 
man, who was treated in the United States Naval Hospital at 
Canacao. This patient had contracted his infection during a stay 
in Guam, where he had become intimately associated with natives 
in whose families Gangosa had been known. 

A further case with complete post-mortem findings was 
described by Musgrave and Marshall (1907) in a native of Santo 
Domingo de Basco, Batan Islands, being the first case reported 
from the Philippines. Careful examination of the blood was made 
by these authors. According to their findings the blood did not 
show any marked pathological changes; the differential count 
proved that the relative numbers of the various types of leucocytes 
were normal. 

In 1909, N. T. McLean observed two patients suffering 
from what he believed to be Gangosa on the occasion of a visit 



214 


to the Hospital at Gonaives, Haiti, and stated that according to 
information that he collected numbers of such cases were supposed 
to have occurred at Port au Prince. 

A case was also published by Stitt in a young Filipino who had 
never been outside Cavite Province (Philippine Islands). 

In 1910 Garrison pointed out that there were 327 cases of 
Gangosa in Guam, and pronounced the opinion that according to 
his experiences from a clinical, and particularly from a 
therapeutical standpoint, Gangosa was to be considered a very 
late form of Syphilis—perhaps a fourth stage. Odell (1911) reported 
a positive Wassermann reaction in 82 % of his cases. Anti¬ 
syphilitic treatment seemed to benefit all cases considerably. 

Rossiter (1912) described a disease in a two year old child where 
the destruction of the hard and soft palate were the most prominent 
lesions. In specimens taken from the ulcerated surfaces numerous 
Spirochaeta per tenuis were found. Two months previously the 
child had suffered from typical framboesia. 

Also in 1912 Ziemann drew attention to the fact that a disease 
resembling Rhinopharyngitis mutilans occurs not infrequently 
amongst the negroes of Kamerun. 

I observed cases of Gangosa for the first time in the Torres 
Straits Islands, whilst on a short journey during the latter end of 
1910. 

Then I was inclined to consider this disease either as a peculiar 
manifestation of syphilis or as a hitherto undescribed disease 
occuring on Murray Island. Elkington referred to most of these 
cases in the yearly report of 1912, adding a number of cases which 
he discovered on Darnley, Boigu, Saibai and Dauren Islands. The 
disease must have been prevalent on Murray Island for nearly a 
century, as it is, according to Elkington, referred to by Dr. Wilson, 
R.N., in an account of a visit made by him to Murray Island in 
1822. 

The occurrence of cases of Gangosa in New Guinea has been 
noted previously, but these cases had been regarded by Sir William 
Macgregor as lupus. 

During my short journey in New Guinea typical cases of 
Gangosa were seen in villages on the south coast, especially in 
Kerapuna, a fairly large village, where as many as ten cases were 



215 


examined. Furthermore a number of natives suffering from 
Gangosa were encountered in the western parts of British New 
Guinea and in the Mekeo District, and there the earliest stages of 
the disease were seen, showing peculiar manifestations which did 
not altogether coincide with the description given by previous 
observers elsewhere. 


CLINICAL ACCOUNT OF THE DISEASE 

According to Castellani and Chalmers the disease ‘ begins as a 
sore throat, when on examination a nodule can be seen on the 
back of the pharynx, the posterior pillars of the fauces or the 
edge of the soft palate. This ulcer usually spreads rapidly at first, 
but more slowly later, and destroys the soft palate first, later the 
bony parts of the palate and the nasal septum. In consequence the 
skin of the nose falls in, and the cavity of the nose and mouth are 
connected. 

1 In consequence of the destruction of the soft and hard palates 
the ulceration may extend afterwards to the face or lips or affect 
the pharynx. As a rule a pungent odour is given off, whilst a 
slight discharge of granular and necrotic debris is present. The 
ulceration may progress continuously for a period of 10-35 years, 
or it may advance at certain times and be quiescent at others, or it 
may cease at any time, leaving a chronic ulcer.* 

So far as observed, the cases of Gangosa met with in New 
Guinea may be conveniently divided into three groups. 

Firstly. Early cases in which the patients had been ill for a 
few months only and where a small area of the skin of the face was 
found to be affected. The cartilaginous septum had been destroyed 
but the disease had not caused any extensive destruction of the 
bone. 

Secondly . Cases in which the ulceration had advanced 
further. Large areas of the face were found to be the seat of 
extensive weeping sores. The bone near the affected parts had 
become implicated, and the disease had led to the destruction of 
the skeleton of the nose and the hard palate. 

Thirdly. Cases in which the active ulcerating process had come 
to a standstill. Extensive cicatrisation had taken place at the 



21 6 


seat of the ulceration, as a rule the nose had disappeared, and was 
represented by a smaller or larger opening. In one extreme case 
the opening was so small that the woman in question was able to 
smoke comfortably a cigarette through the small opening—the only 
remains of the nose. In another case the whole of the back of 
the pharynx could be seen without any difficulty through the large 
opening in the middle of the face. 

The uncertain etiology of Gangosa and its similarity to tertiary 
syphilis in its clinical manifestations makes it advisable to refer 
in greater detail to the history of a few of the typical cases met with. 


EARLY CASES 


Only a comparatively small number of early cases were seen. 

Case i. (PI. XIV, fig. i.) Apau Kaianga was an unmarried girl of about 
12 years of age. Her parents as well as four brothers and sisters were alive and 
well. Her disease began about one month previous to my visit as a sore below 
her nose which spread rapidly, affecting especially the septum, the left ala of the 
nose, the upper lip and the left corner of the mouth, and had already destroyed 
part of the cartilaginous septum. 

The examination revealed the nose considerably flattened, and also thickened on 
account of oedematous swelling. The left ala was the seat of a large raised dry ulcer, 
covered by a thin scab, below which there was a reddish uneven granulating surface. 
This ulceration had extended to the upper lip, being confined to its central part, 
just below the nostrils. The left corner of the mouth was the seat of a similar 
ulcer which showed an uneven surface. The upper lip around the ulceration 
was swollen and oedematous and its surface was intersected by purplish irregular 
raised lines simulating scar tissue. The cartilaginous septum had been partly 
destroyed by the ulceration, whereas the osseous septum .was still intact. The 
hard and soft palate were of normal appearance. 

Case 2 (PI. XIV, fig. 2.) Ame Aua was a woman about 25 years of age, 
belonging to the same village. Her father was still living, her mother had died. 
Some time ago she was married, her three children were alive and well, except 
for manifestations of Yaws. 

According to her own statement she had been ill for four months only, the 
disease nn her had begun as a small sore on the tip of her nose which had spread 
gradually into the nasal cavity. The examination revealed a fairly large ulcer 
on the nose, a yellowish greasy scab covering the raw granulating surface. The 
cartilaginous septum of the nose had been destroyed by ulceration and the nose 
was, in consequence, flattened. A similar ulcer below the nostrils extended into 
the nasal cavity. On the right cheek was a narrow raised line resembling oedematous 
scar tissue. The hard and the soft palate and the pharynx did not show any 
pathological changes. 



2I 7 


Case 3. (PI. XIV, fig. 3.) The woman belonged to Kerapuna, a large village 
on the south-east coast of New Guinea. She had been married and was the mother 
of six children. Four of them were healthy and two were in the secondary stage 
of Yaws. 

Her upper lip showed the same scar formation as described in the two previous 
cases, the scars being oedematous and raised in the centre. The free edge of the 
skin of both the alae showed ulceration and both nostrils were closed up by yellowish 
scabs. 

Case 4. (PI. XIV, fig. 4.) Ame Kwipa, a young girl, belonged to Veipa, a 
large village in the Mekeo District. She had had small sores on her face, especially 
on her right cheek for about one year, and these had shown a tendency to heal 
during the last two months. Her brothers and sisters were healthy. 

When examined her nose appeared broadened, its skin swollen and oedematous. 
The upper lip, especially the right side, was swollen, oedematous, and showed 
raised irregular lines resembling scars. The right cheek was the seat of a 
number of small pustules which contained whitish pus. 

Case 5. (PI. XV, fig. 5.) Ainye, a girl about 14 years of age, belonged to 
Bebeo, a small village in the Mekeo District. Her father was alive and well, 
her mother and one brother had died. Her illness commenced about one year 
previously with a small sore on the upper lip below the nose, which had spread 
slowly and gradually to the nose and downwards to the mouth. 

When examined her nose was flattened, showing an uneven surface; the mucous 
membrane of her nostrils was covered by a yellowish scab. The upper lip was 
swollen and succulent in appearance, whilst the left angle of her mouth was drawn 
up by the formation of scar tissue. Her body was clean. 

Case 6. (PI. XV, fig. 6.) The patient was a young unmarried girl, Wabnagi, 
belonging to Kaile, a village on the south-east coast of New Guinea. Her mother 
was alive, her father dead. Her disease began about two years previously as a 
small sore on the upper lip just below the nose, which had spread gradually to the 
cheeks and into the interior of her nose. 

At the time of examination her nose was flattened and had sunk in, as the 
cartilaginous septum had completely disappeared. The surface of the nose was 
uneven and rough ; the right half of the upper lip was drawn up on account of 
extensive scar formation. The same scar formation was seen on both cheeks, 
more extensively, however, on the right side. The skin of the affected parts 
was very thin and shiny, intersected by raised streaks of apparently dense scar 
tissue.. Here and there on the nose and cheeks were small ulcerations covered 
by a yellowish scab. The hard and soft palate did not show any pathological 
changes. 

Case 7. (Pl. XV, fig. 7.) Was a young unmarried woman belonging to 
Kerapuna. Her parents as well as two younger sisters were alive and well. Her 
disease began when a small child with sores on her face which had since spread, 
implicating the nose. She showed large and extensive ulceration on her face ; 
the nose had completely disappeared and was represented by a small opening 
in the middle of the face surrounded by irregular ulcers covered by yellowish 
scabs. The upper lip was partly destroyed by extensive ulceration. A large 
ulcer on her left cheek had a raised granulating surface and irregular edges, secreting 
copiously a slightly yellowish clear, foul-smelling, discharge. The left cheek 
showed scar formation. The hard and soft palate were normal. 



218 


LATER CASES 

Case 8. The patient, a man about 22 years of age, belonged to Hula, a 
village on the south-east coast of New Guinea. His mother had died, his father 
was alive and healthy. He was married and had three normal children. According 
to information the disease had commenced about five years previously with a sore 
on the forehead over his left eye-brow, which had spread gradually until the 
greater part of his face was affected. His face, when examined, was badly disfigured; 
the nose was flattened and was sunk in. Deep scar formation on the right side 
of his forehead caused the upper eyelid to be drawn up. The skin on the left 
side of his forehead showed extensive ulceration which had spread downwards 
and caused the destruction of his left eye. The central part of Jiis upper lip was 
partly destroyed. The hard palate was perforated, showing a round hole of 
about 2 cm. in diameter, with smooth rounded edges. 

Case 9. Was a woman, Kila Alukma, of about 40 years of age, living in the 
same village as the previous case. Her parents were dead, her two daughters 
were married and apparently well. 

Her disease had begun about ten years previously ‘ with a pain in the stomach 
which gradually worked up to her mouth and came out through her nose/ She 
pointed out that there had been a sore in her mouth before her face became 
affected. This case was very similar in appearance to Case 8. There was extensive 
scar formation on the face round the nose, the upper lip was drawn up on account 
of dense scar tissue. A stinking discharge from the nostrils was noticed; the 
hard palate was perforated, the oral and nasal cavity communicating through a 
round opening of about 3 cm. in diameter. 

Case 10. Was an unmarried woman of about 35 years of age belonging 
to Kaile on the south-east coast of New Guinea. Her mother was still alive, 
her father was dead. About ten years previously a small ulcer had started on 
her nose and soon began to spread. At the same time ulcers appeared on her legs. 

When examined the cartilaginous septum of her nose had disappeared and 
extensive scar tissue formation was observed around the nose, on both cheeks 
and on the upper lip, the mouth and eyelids being normal. The hard palate 
was perforated. Her legs, as well as her left forearm were the seat of deep ulcers 
of varying size some of which extended as far as the bone, whilst some of them 
were covered by a greyish greasy scab. 

Case i i. (PI. XV, fig. 8.) Was a man, Aiaba, about 40 years of age belonging 
to Kivori village. Both his parents were dead. He was married and the father 
of six healthy children. His illness had begun about four years previously with 
a small sore on the upper lip just below the nostrils, which spread at first into the 
nasal cavity and later to his face. He had lost the toes of his right foot in an 
accident when he was a small boy. 

The patient’s face was of horrible appearance. The nose had completely 
disappeared, leaving a large hole in his face through which the posterior wall of 
the pharynx could be seen. A large irregular granulating sore on his forehead 
was covered by a yellowish dry scab under which thick pus had collected. A similar 
sore was on his right cheek showing here and there recent scar formation. The 
upper lip was the seat of extensive scar formation. The skin of his left cheek 
was swollen and oedematous. 

The hard palate was perforated and the oral cavity communicated with the 
nasal cavity by means of a large round opening with sharp edges. 



219 

THIRD STAGE OF THE DISEASE 


Case 12. (PI. XVI, fig. 9.) A woman, Ugre, about 25 years of age, who 
lived in Kerapuna, a village on the south-east coast of New Guinea. She had 
been ill, according to information, for about ten years; her nose had been affected 
long before her mouth became involved. When she fell ill her husband married 
a second wife. Both were seen and were in excellent health. When examined 
the skeleton of her nose had disappeared and there was a hole in the centre of her 
face, the surrounding skin as well as the skin of the upper lip and forehead showed 
extensive scar formation, indicating that the whole face had been previously 
affected by the disease. 

Case 13. (Pl. XVI, fig. 10.) An unmarried woman, about 32 years of age, 
was seen in the same village as the previous case. Her father had died, her mother 
was still alive and in good health. Her disease had started when she 
was quite a small child. When examined the bony skeleton of her nose had 
completely disappeared, the nose was sunk in so that the nostrils were only 
represented by a small opening. Both the upper and the lower lips had disappeared, 
the skin of the face directly joining on to the gum, so that the teeth were 
unprotected; the skin of her face was one mass of scar tissue, the right lower 
eyelid was drawn down by scars giving rise to an ectropion. The left side of 
her throat and her left shoulder were of the same appearance as the skin of her 
face, showing extensive scar formation. Her soft and hard palate did not show 
any pathological change. 

Case 14. (PI. XVI, fig. 11.) An unmarried woman of about 35 years of age. 
She had been ill for quite a long time, in all probability between ten to fifteen 
years. Both her parents were alive and in good health. Her disease began on 
the ‘ outside of her nose 9 in the form of a small sore which had spread slowly 
implicating the lips and mouth. Some time ago the ulcerated parts had begun 
to heal up. Her face was completely disfigured. The whole outer nose had 
disappeared as well as the anterior part of her hard palate, so that the oral and 
nasal cavities communicated extensively. The posterior part of the hard and 
soft palate were still intact. The central part of the upper lip and the lower 
lip had disappeared, the outer skin joining immediately on to the mucous membrane 
of the gum. There was extensive scar formation on the skin of her face and on the 
neck which closely resembled the scars after deep burns. 

Case 15. A woman of about 35 years of age belonged to Ififu (Mekeo 
District). Her parents were dead, two brothers were alive and well, one of them 
showing scar formation around the mouth, most probably the remainder of Yaws. 
She was married and mother of a healthy girl. Her illness began when she was 
a girl, in the form of a small sore on the upper lip. Subsequent to the infection 
of the face a large ulcerating sore developed on the abdomen, around the navel 
which, however, soon healed up. Her mouth was small and deformed, the left 
angle drawn up by scar tissue. Extensive scar tissue formation was seen on the 
upper lip and both cheeks, the skin otherwise being smooth, shiny and very thin. 

Case 16. (PI. XVI, fig. 12.) Was an unmarried woman, Kaiyo, about 35 years 
of age. Her parents were dead. The disease began at a very early age with 
a sore on the left foot soon after she had begun to walk. Later on a small ulcer 
appeared on her upper lip below the nostrils which gradually spread, eating away 
the whole upper lip and the nose. At the time of examination, her nose as well 
as her upper lip had completely disappeared, a small round hole representing 



2 20 

the nasal opening. Extensive scar formation was noticeable around the mouth 
and on the cheeks. Her left eye had been destroyed by the disease. The hard 
and soft palate did not show any pathological changes. 

Case 17. Was a middle-aged woman. Her parents were dead. She had 
four brothers and sisters, two of whom had died. She had never been married. 
Her disease began some time ago with an ulceration inside her nostrils. Her 
nose had been destroyed by the disease, and the back of her pharynx was visible 
through a large triangular opening in the face. The anterior part of the hard 
palate was destroyed, but the posterior part was still existent. Her mouth was 
deformed, the right angle drawn up by scar tissue. 

The case histories recorded above describing a few typical cases 
out of the many examined, prove that Gangosa, or Rhino¬ 
pharyngitis mutilans, is a definite morbid entity, and can be 
differentiated from other diseases causing similar lesions, such as 
Syphilis, Lupus and Leprosy. It is a very chronic but very rarely 
fatal complaint. A great number of the patients recover without 
specific treatment, mostly, however, after the morbid process has 
brought about extensive destruction of the face. 

From the number of cases observed in the different stages, 
Gangosa in New Guinea seems to differ in its earliest stages from 
the disease described by Castellani and Chalmers. 

According to my experience in New Guinea, Gangosa usually 
begins as a small ulcer on the upper lip, just below the nose. Only in 
a small number of cases did patients state definitely that the palate 
was affected before any other part of the face. This small ulcer 
soon begins to spread, destroying at first the fleshy parts of the face, 
such as the lips, the alae and the skin of the cheeks surrounding 
the nose. As a rule, the tissue in the near neighbourhood of the 
ulcer is swollen on account of the oedematous infiltration, the surface 
skin is glossy and intersected by reddish, raised lines, resembling 
in some respects a relief map of mountains. 

In the earliest stages the ulcer may spread gradually or some¬ 
times rapidly, finally implicating the bones of the infected regions, 
destroying at first the cartilaginous parts of the nose, and later on 
the bony skeleton and the soft and hard palate. 

The ulcers are raised, possess irregular edges, and are not well 
defined from the surrounding tissues. The surface of the ulcer is 
formed of granulating tissue, which secretes a malodorous yellowish 
discharge, and is often covered by a dark yellowish scab under¬ 
neath which pus accumulates. Very soon, however, some of the 



221 


ulcers show a tendency towards healing. The granulation tissue 
begins to discharge less, and after a varying period new smooth 
a skin grows over the ulcer from the surroundings, often, however, 
"breaking down again, and giving rise to renewed ulceration. 

Some time afterwards dense scar tissue is formed, leading to 
the deformities of the face so characteristic of the disease. 

Now and again only a single primary ulcer is seen, at other 
times a number of ulcers appear simultaneously. 

The morbid process may come to a standstill at any stage of the 
disease, even before it has led to extensive destruction. 

The tendency to scar formation is a marked feature of the 
disease, and in this respect Gangosa simulates Syphilis, so that in 
some of the late cases a differential diagnosis would be practically 
impossible. 

Some of the patients who showed typical symptoms of Gangosa 
in their faces were suffering from large ulcerations on other parts 
of the body, especially on the legs. It is impossible to decide 
whether these ulcers corresponded' only to Ulcus tropicum, or were 
due to the same parasites as the face lesions. 

It is interesting to note that the great majority of cases seen 
were women, only a comparatively small number of men being 
affected. 


THE ETIOLOGY OF THE DISEASE 

As the etiology of the disease is unknown, material was collected 
for microscopic examination. Special attention was paid to the 
early lesions with a view of finding a possible parasite in the 
oedema fluid. It may well be understood that the examination of 
the secretion of open sores of advanced cases did not seem to offer 
much opportunity for the forming of a definite opinion; smears 
taken from the secretion of open wounds always contain enormous 
numbers of bacteria and spirochaetes of varying shapes and sizes, 
which are to be considered as secondary infections. 

From the earliest cases showing the marked oedematous swelling 
of the affected parts as described above, oedema fluid was obtained 
by pricking the skin over the lesion with a needle in the same way 
as serum would be obtained from a leprous nodule. The specimens 



222 


were stained with Giemsa's stain. In the oedema fluid of five early 
and two later cases, in addition to a small number of red and white 
corpuscles, small bodies were found resembling yeast cells. The 
majority of them occurred free, only a small number was contained 
in leucocytes (PI. XVII, figs. I and 2). These cells showed a marked 
polymorphism; some of them were round, others oval or pear- 
shaped (comp. figs. 3 and 4), and were seen either singly or in 
groups. 

The cells measured from 075 fi to 4 /*, and consisted of a 
lightly bluish staining cytoplasm showing a typical network 
structure, a great number of them contained a distinct and 
sharply-defined vacuole (PI. XVII, figs. 5-7) and small roundish 
masses which resembled chromatin in their staining reaction. These 
chromatin-staining masses occurred either singly, being more or less 
sharply defined, roundish or oblong in shape (figs. 7, 8, 13) or as 
number of similar bodies enclosed within the cell (figs. 9, 10, 

14. 15)- 

Budding of the cells in its various stages could be frequently 
observed. 

In the earliest stages only a small protuberance on the periphery 
of the cell was seen, consisting of cytoplasm only (figs. 3 a, 6, 14). 

In the more advanced forms this protuberance had increased in 
size, and at this stage a small chromatin-like staining mass was 
often seen at the base of the bud (figs. 3 b, 16). Sometimes the bud 
itself contained one or more chromatin-like staining particles 
(figs. 7, 13). 

The bud increased in size, and usually became detached before 
it reached the same size as the parent cell (figs. 9, 10, 16), and in 
this case invariably contained chromatin-like staining particles and 
often a distinct unstained vacuolic area (fig. 18). 

In many instances only one bud was formed, not rarely, how¬ 
ever, two, three or even more buds were seen still connected with 
the cell (figs. 7, 15). 

Forms as seen in fig. 17 were noticed only very rarely. In these 
forms the cell consisted of a central portion stained lightly reddish by 
Giemsa's method, containing one or more distinct chromatin-like 
particles, the whole being surrounded by a thick bluish pellicle. 

From their morphological appearance there is no doubt that the 



223 


cells described are to be considered as parasites belonging to 
the genus Cryptococcus of the family Saccharomycetes. The 
characteristic features of Cryptococcus is the reproduction by 
budding only and the absence of endospores and ascospores. 

In these parasites only multiplication by budding was observed, 
endospores have never been seen, although carefully sought. 

The parasite itself has all the typical features of yeast cells. 
Unfortunately circumstances only permitted of the collecting of 
dry films, which were stained by Giemsa’s method, and nothing 
definite can be said about the nuclear details. When stained by 
Gram’s method the cells did not become decolourised. 

Parasites of the genus Cryptococcus are well known to produce 
granulating sores in man and beast; the best known and most 
studied disease due to Cryptococcus is probably epizootic 
lymphangitis in horses, occurring in different parts of the world. 

Gilchrist and Stokes (1898) describe a similar parasite in man 
from a case of chronic ulcerative dermatitis, resembling Lupus, under 
the specific name of Cryptococcus dermatitis. Busse (1894) isolated 
Cryptococcus hominis from a purulent periostitis of the tibia of a 
woman. 

On the other hand, yeast cells occur very frequently in open 
sores of every description. 

These parasites were found in great numbers in the oedema 
fluid of the early cases of Gangosa, where open sores had not 
formed. Moreover, specimens of the oedema fluid did not contain 
any other microorganisms besides the yeast cells and a small 
number of red and white blood corpuscles. 

In the seven cases from which oedema fluid could be obtained 
these parasites were found, most numerously in Case 11, where early 
and late manifestations were present at the same time, and in which 
the disease seemed to be progressing most rapidly. 

The finding of a Cryptococcus in seven cases out of eight 
examined, suffering from Rhinopharyngitis mutilans, justifies the 
conclusion that one may regard this parasite as the cause of the 
disease in question. 

The fact that blastomycetes are known to cause similar skin 
lesions in man and animals is an additional argument in favour 
of the Cryptococcus being the etiological agent of Gangosa. Wc 



224 


therefore propose for this parasite the specific name of Cryptococcus 
mutilans. 

Unfortunately, circumstances did not allow attempts at cultiva¬ 
tion. Animal experiments have been performed at our instigation 
by Dr. Elkington whilst visiting Murray Island. Two monkeys 
were inoculated on the eyelid and intra-nasally with the secretion 
of open sores, but in spite of prolonged observation none of the 
animals in question showed any lesions whatsoever. 


CONCLUSION 

Rhinopharyngitis mutilans is a disease which occurs not 
infrequently amongst the native population of the south coast of 
British New Guinea, and is a blastomycosis due to Cryptococcus 
mutilans , n.sp. 


REFERENCES 

Breinl (1910). Report of the Australian Institute of Tropical Medicine for the year. 

Busse (1894). Ueber parasitare Zelleinschlusse und ihre Ziichtung. Ccntralbl. f. Bakt. 
Bd. XVI. 

Castellani and Chalmers (1910). Manual of Tropical Medicine. 

Elkington (1912). Annual Report of the Commissioner of Public Health, Brisbane. 

Fordyce and Arnold (1906). Journal of Contagious Diseases , Vol. XXIV, 1. 

Garrison (1910). Gangosa in Guam. Bulletin of the Manila Medical Society , Vol. II, 
No. 10. 

Gilchrist and Stokes (1898). The presence of an oidium in the tissue of a case of 
Pseudolupus vulgaris. J. Hopkins Hosp. Rep ., Vol. VIII, No. 64. 

Leys (1906). Journal of Tropical Medicine , Vol. IX, p. 47. 

McLean (1909). Gangosa in Haiti. United States Naval Medical Bulletin , Vol. Ill, p. 141. 

Mink and McLean (1906). Journal of American Medical Association , Vol. XLVII, 
p. 1166. 

Musgrave and Marshall (1907). Gangosa in the Philippine Islands. Philippine 
Journal of Science, Vol. II, p. 387. 

Odell (1911). Gangosa a form of Syphilis. United States Naval Medical Bulletin, Vol. V, 
No. 4. 

Rossiter (1912). Report of a case resembling Gangosa in which Treponema pertenuis 
was present. United States Naval Medical Bulletin , Vol. VI. No. 1. 

Stitt (1907). A case of Gangosa in a white man. United States Naval Medical 
Bulletin , Vol. I, p. 96. 

-11910). A case clinically resembling Rhinopharyngitis mutilans. United States Natal 

Medical Bulletin , Vol. IV, No. 4, p. 524. 

Wilson (1822). Voyage round the World. 

Ziemann (1912). Discussions during the Meeting of the German Tropical Society. Archiv f. 
Schiffs - u. Tropenbygiene, Bd. XVI, Beiheft, p. 85. 




226 


EXPLANATION OF PLATES 

Plate XIV 


Figs. 1-4. Cases of Gangosa (Rhinopharyngitis mutilans). 




.-{finals Trop. Med. 







228 


Plate XV 


Figs. 5-8. Cases of Gangosa (Rhinopharyngitis mutilans) 



A finals Prop. Med. & ParasitolVol. IX 


PLATE XV 



C. Ttitling «S^ Co., Ltd., Imp 







130 


Plate XVI 


Figs. 9-12. Cases of Gangosa (Rhinopharyngitis mutilans). 






Plate XVII 


All the figures were drawn with a large Abbe drawing apparatus, 
using 2 mm. apochromatic oil immersion. 

For Figs. 1-2. Compensation ocular 8 was used. 

For Figs. 3-18. Compensation ocular 18. 

The specimens were fixed in alcohol and stained by Giemsa’s 
method. 

Figs. 1-2. Intracellular forms of Cryptococcus mutilans. 

Figs. 3-4. Groups of parasites. 

Figs. 5-7, 9, 10, 12-16, 18. Different stages of budding. 

Figs. 8, 11. Large forms. 

Fig. 17. Apparently encysted form, rarely seen. 




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CRYPTOCOCCUS MUTILANS. 




233 


THEILERIA TACHTGLOSSI (n.sp.) 
A BLOOD PARASITE OF TACHT- 
GLOSS US ACULEATUS 


BY 

HENRY PRIESTLEY, B.Sc., M.B., Ch.M. 

FROM THE AUSTRALIAN INSTITUTE OF TROPICAL MEDICINE 


(Received for publication 20 May y 1914) 
Plate XVIII 


Hitherto blood parasites of Monotremes have not been 
described. 

Gilruth, Sweet and Dodd (1911) refer to the occurrence of 
Attaplasma marginalc (Theiler) in the blood of Echidnas, but 
according to their conception the presence of these bodies has no 
pathological significance. 

The blood of an Echidna, from the neighbourhood of 
Townsville,* on examination, showed the presence, within the red 
blood corpuscles, of numerous parasites resembling the Thcileria of 
East Coast Fever. On rare occasions the same structures, referred 
to by Gilruth, Sweet and Dodd as Anaplasma } could be seen. 

When the blood of the animal was again examined after an 
interval of a week, most of the parasites had disappeared from the 
peripheral circulation, and so the animal was killed and films made 
from the blood and different organs. 

Dry films were fixed in alcohol and stained with Giemsa’s 
solution. It was found practically impossible to obtain satisfactory 
results with wet fixing and staining methods for the blood films. On 
the other hand the detailed structure of Koch’s bodies could be 
studied to advantage in organ films after wet fixation in sublimate 
alcohol and staining by Giemsa’s method. 

* I take this opportunity of thanking Mr. J. Humphrey of the Haughton River through 
whose kindness the animal was obtained. 



234 


MORPHOLOGY OF THE PARASITES IN THE PERIPHERAL 

BLOOD 

In their morphological appearance the parasites resembled 
closely Thcileria parva of East Coast Fever. 

They occurred, invariably, within the red blood corpuscles 
without causing any apparent alteration in the shape and form of 
the cells. 

Bacilliform, ovoid, pyriform, round and comma-shaped 
parasites were seen, but on careful examination intermediate forms 
were often encountered. Typical examples of each class could, 
however, be easily distinguished. 

The bacilliform parasites (PI. XVIII, figs. 1-3) occurred most 
frequently. They measured i‘S yu to 3 o yu by 0*3 M to O'8/x, and 
were cither perfectly straight or slightly curved. 

The cytoplasm stained dark blue and the chromatin was 
situated either in the centre or at one end of the parasite. 

The comma-shaped parasites (fig. 5) measured 2 0 yu to 3 0 n 
by 0 8 yu to 1*2 yu, were broader and were rounded off at one end. 
The cytoplasm stained less deeply and an unstained vacuolar area 
was frequently present. 

The ovoid and pyriform parasites (figs. 4 and 6) measured 1*0 yu 
to 3*0 ft by o‘5 yu to 1*2 yu, and no sharp line of distinction could 
be drawn between these two types. A vacuole was almost always 
present, with the chromatin, as a rule, in close proximity to this 
unstained area. Sometimes, however, the chromatin occurred in 
the form of an irregular band across the centre of the parasite. 

Rounded forms (fig. 7) were the least frequently encountered. 
They measured ro yu to 2*2 yu in diameter, and resembled the ring 
forms of malarial parasites. The cytoplasm, which stained light 
blue, appeared to be condensed peripherally, surrounding a central 
clear area. The chromatin occurred as an oval or round mass, or 
was crescentic in shape and situated at tne periphery. Sometimes 
two masses of chromatin were seen in the same parasite. 

Multiple infection of red blood corpuscles was very rare. In 
five slides three corpuscles were found containing two parasites, 
one contained three parasites and one contained four parasites 
(fig- 9 )- 

The so-called cross forms with chromatin particles at each pole 
were seen on two occasions only (fig. 8). 



2 35 


As in infections with Theileria parva , the so-called Koch’s 
bodies occurred in small numbers in the peripheral blood, but in 
great numbers in smears made from the organs, particularly the 
liver, spleen and lungs. They were either free or included in 
leucocytes or endothelial cells, and represent in all probability an 
asexual multiplication of the parasite. 

The smallest forms within the leucocytes somewhat resembled 
cell granules. They were about i fi in diameter, rounded, with 
pale blue cytoplasm and a comparatively large chromatin mass in 
the centre. The largest forms occurring intracellular or free in 
about the same proportion were rounded or oval in shape, 
measuring up to 12 n in diameter. 

The cytoplasm stained by Giemsa’s solution was either dark or 
light blue, showing the typical network structure; the chromatin 
occurred in the form of numerous granules, irregular in shape and 
size and unevenly distributed throughout the parasite. In many 
instances a great number of chromatin granules were aggregated 
into masses forming irregular bands across the cytoplasm, at the 
same time chromatin particles were dotted over the remainder of 
the parasite (fig. 16). 

All intermediate stages in the leucocytes were observed between 
the smallest forms containing only one small chromatin granule 
and little cytoplasm and the largest forms with a great number of 
chromatin granules. The cytoplasm of the largest intracellular 
forms was usually well defined from the cytoplasm of the host 
cell; in some cases, however, no such differentiation could be made 
out, and in these instances irregular chromatin granules were 
unevenly distributed throughout the host cell. It would seem that, 
in these cases, the parasite had ruptured, leaving the chromatin 
granules, most probably surrounded by a very small amount of 
cytoplasm, free within the host cell. The fact that, now and 
again, fully developed parasites, of the same type as those found 
in the red corpuscles, were seen in leucocytes in addition to 
chromatin granules, suggests that the majority of the chromatin 
granules may develop into the small blood forms. 

As in the case of Theileria parva , when the leucocyte contains 
a large Koch’s body the cell nucleus may be disintegrated. 

The parasite of the Echidna closely resembles, in its life 
history, Theileria parva as described by Gonder. 



It may be assumed from these observations that the chromatin 
of the small forms which have entered leucocytes divides with a 
corresponding increase in the amount of cytoplasm. 

Finally the larger forms burst, setting free small parasites 
resembling closely the forms found in the red corpuscles. It seems 
obvious, then, that the parasite undergoes a definite schizogony 
within the leucocytes. 

A gamogonous and an agamogonous generation, as described 
by Gonder for Theileria parva , could not be followed out in the 
parasite of the Echidna. 

The occurrence of bacilliform parasites, cross forms and Koch’s 
bodies places the parasite of the Echidna in the genus Theileria , and 
the specific name of Theileria tachyglossi is proposed. 

The Echidna from which the parasites were obtained was 
infested with ticks— Aponomma dec or o sum (L. Koch)—and was 
apparently healthy. 


SUMMARY 

A new species of the genus Theileria was found in an Echidna, 
Tachyglossus aculeatus . 

This parasite, for which the name Theileria tachyglossi is 
proposed, resembles very closely Theileria parva. Koch's bodies 
were found in great numbers in the internal organs, and to a certain 
extent in the peripheral blood, and represent a stage in the life 
history of Theileria tachyglossi in the warm-blooded animal. 


REFERENCES 


Gilruth, Sweet and Dodd (1911). Observation) on the occurrence in the blood of vaiious 
animals (chiefly Monotremes and Marsupials) of bodies apparently identical with 
Ana pi <um a marginal Theiler, 1910. Parasitology Vol. IV, pp. 1-6, 1 plate. 

Gonder (1909-10). Report of Government Veterinary Bacteriologist. Transvaal 1909-10. 
P . 69. 




238 


EXPLANATION OF PLATE XVIII 


All the figures were drawn by means of an Abbe drawing 
apparatus from preparations fixed dry and stained with Giemsa’s 
solution. Zeiss apochr. homog. 2 mm., comp, ocular 18 except 
figs. 15 and 16, in which comp, ocular 8 was used. 

Figs. 1-3. Bacilliform parasites. 

Fig. 4. Ovoid parasites. 

Fig. 5. Comma-shaped parasite. 

Fig. 6. Pyriform parasite. 

Fig. 7. Round parasite. 

Fig. 8. Cross-form parasite. 

Fig. 9. Four parasites in one corpuscle. 

Fig. 10. Anaplasma. 

Fig. 11. Koch’s body, large free form with deep-staining cyto¬ 
plasm. 

Fig. 12. Koch’s body with pale-staining cytoplasm. 

Figs. 13, 14. Free parasites in leucocytes. 

Figs. 15, 16. Intracellular Koch’s bodies. 





THEILERIA TACHYG LOSSI. 




239 


AN INVESTIGATION INTO THE CAUSES 
OF THE PREVALENCE OF ENTERIC 
FEVER IN KINGSTON, JAMAICA ; WITH 
SPECIAL REFERENCE TO THE QUESTION 
OF UNRECOGNISED CARRIERS 


BY 

H. HAROLD SCOTT, M.D. (Lond.), D.P.H. (Hons.) 

GOVERNMENT BACTERIOLOGIST ; PATHOLOGIST TO THE PUBLIC GENERAL HOSPITAL, KINGSTON, JAMAICA 


{Received for publication I February , 1915) 


Ten Charts 


CONTENTS 


I. Introductory . 

II. Water Supplies . 

III. Milk and other forms of Food . 

IV. Flies. 

V. Sewage Disposal . 

VI. Carriers . 

VII. Bacteriological Investigations :— 

i. Method employed . 

ii. Cases with post-mortem signs of enteric fever giving positive results 

on cultivation of the bile. 

iii. Cases giving similar results, but showing no evidence post-mortem of 

enteric fever. 

iv. Deductions from the fact of isolation of the Bacillus typhosus from 

the gall-bladder or its contents . 

v. Peculiarities of the organism isolated in some instances . 

vi. Length of stay of the organism in the gall-bladder . 

vii. Presence of bacilli with and without obvious inflammatory changes 

in the gall-bladder. 

viii. The question of atypical cases without characteristic changes 

VIII. Personal Contact. 

IX. Legislative Recommendations . 

References. 

Table . 

Charts 


page 

240 

2 45 

2+7 

248 

249 
251 


255 

256 
158 

261 

262 
265 

265 

266 

267 

268 


27^ 









240 


I. INTRODUCTORY 

The^extensive prevalence of enteric fever in the town of Kingston, 
Jamaica, and its suburbs has been for some years a cause 
of considerable uneasiness, amounting at times to actual alarm. 
This uneasiness is fully justified when it is considered that the city 
is ideally situated in that it is exposed to bright sunshine practically 
all day long and on nearly every day of the year, for the weather 
might be regarded as that of perpetual spring and summer with 
occasional showers. The disposition of the town is that of a gentle 
slope from the upper parts right down to the sea, and therefore with 
an ideal natural drainage. The obvious inference is that the reason 
of the undue prevalence of enteric fever must be looked for in the 
inhabitants themselves. Part of the town is provided with a water- 
carriage system of sewage disposal, but a large part is not. This 
part reminds one very much of the old time military camp, when 
each fresh company of troops arriving, encamped upon or close by the 
latrines of the last, or, when the troops were stationed in a place for 
some time (comparable to the stationary population of Kingston), 
they were from a sanitation point of view living on their own 
dunghill. 

Chart I, compiled from figures which have been given to 
me by the kindness of the Acting Medical Officer of Health, 
shows that the number of notifications of typhoid fever begins to 
increase in March and reaches a maximum in June. A considerable 
proportion of these, of course, are notified by medical practitioners 
in the city without any bacteriological examinations, and con¬ 
sequently include many which are perhaps not enteric fever at all. 
I think this inference is warranted, because if the figures are 
compared with those from the laboratory, where specimens which are 
sent up from suspected cases are examined (see Table I), it will be 
seen that as a rule more than half those examined give negative 
results. We may, therefore, take it that a certain fairly large 
proportion of those notified as typhoid fever are not true cases of 
the disease. I consider that I am further justified in saying that the 
laboratory figures, although based on a smaller total, are more likely 
to be accurate as regards the disease in question than those given by 
the Medical Officer of Health as 4 notifications/ This must not be 



2 4 I 


implied as reflecting in any way on the diagnostic capabilities of the 
medical practitioners here, but it is an undoubted fact that many 
cases which exhibit a continued fever of obscure nature, though not 
enteric, are diagnosed as such when no bacterial tests—Widal, 
culture of the blood, faeces, urine, etc*,—are carried out. Hence, 
although it is probable that few cases of true clinical enteric fever are 
missed, the number notified would show an excess in the actual 
prevalence of the disease in the clinical sense. This remark applies 
generally to all towns. In hospital records, such as I have more 



Chart I.—Notifications of .Enteric Fever cases in Kingston during 1914. 


particularly to deal with, a different state of things may exist. Having 
a pathological laboratory in the centre of the town, any doubtful case 
is subjected to a blood examination, and thereby two errors may be 
rectified. 

Firstly, by this means patients with obscure febrile symptoms, 
but not clinically indicative of enteric fever, might have to be 
reported as such, owing to positive laboratory findings. 

Secondly, suspected cases, which would be reported as enteric 
fever because of this suspicion, are not so notified owing to negative 




242 


results of laboratory tests, or the notification is delayed till other 
symptoms render the diagnosis clear. 

Although these two points are antagonistic, the first tending to 
increase while the second would lead to a diminution of cases 
notified, the latter certainly preponderates, as is shown by the 
appended Table I, where it will be seen that of the specimens sent 
up some 50 per cent, proved to be negative. In other words, about 
one-half of suspected cases are not suffering from enteric fever, that 
is, the notifications, as stated above, are in excess of the actual 
prevalence of the disease. Cases wrongly diagnosed as enteric 
fever will, it is true, erroneously swell the notifications, but if labora¬ 
tory examinations of such are made free (see p. 269), these 
diagnoses can be rectified later, and the Medical Officer of Health 
can make the necessary alterations in his records. 

One of the main dangers of a wrong diagnosis is that such cases 
may then be traced to the operation of some factor in the spread of 
typhoid infection which really has played no part, the case not being 
one of typhoid fever at all. On the other hand, this is a very small 
point in cpmparison with the overlooking of positive cases. In 
other words, the erroneous diagnosis of a non-enteric case as enteric 
is venial compared with the failure to diagnose a true case because 
the clinical manifestations happen to be anomalous, through omitting 
to make use of a laboratory when it is situated centrally, and thus 
allowing a possible carrier to go about unwarned. 

It must be remembered that among the 4 negative ’ cases are 
included some which on subsequent testing yielded positive results, 
since some specimens are sent up too early in the disease (second or 
third day, perhaps), while in others agglutinin formation may be 
delayed. 

The taking of a sample of blood for a Widal test is simple and 
causes practically no pain, but in the interpretation of the result 
there are certain pitfalls. These have been dealt with in more detail 
elsewhere (Scott, 1913), so it will suffice merely to mention the 
following facts: — 

(1) Though a positive result is obtained in the majority of 
cases (95 per cent.) at some time or other of the disease, 
there is, it would appear, a small percentage in whom it 
is not given. Into the reasons put forward for this I need 



243 


Table I.—Showing the numbers of specimens sent to the laboratory month by month 
for Widal’s test during the year 1914 , with the results. 


Month Number Positive Percentage Negative Percentage Doubtful Percentage 



sent 


positive 

1 


negative 


doubtful 

January . 

21 

7 

33 

i ; 

13 

62 

1 

| 5 

February ... 

26 

8 

31 

>7 

«5 

i 

4 

March . 

54 

2 3 

42 

2 9 

54 

, I - 

2 

4 

April . 

68 

2 7 

40 

39 

57 

2 

3 

May . 

"9 

1 

40 

33 

75 

63 

4 

4 

June. 

109 

59 

54 

47 

43 

3 

3 

J“>y . 

62 

22 

35 

39 

6 3 

| 

1 

2 

August . 

65 

22 

__ 

w 

4 - 

40 

61 

3 

5 

September ... 

69 

21 

3° 

46 

66 

3 

4 

" “ 1 _ 1 1 

October . 

7 1 

19 

2 7 

5* 

70 

2 

3 

November ... 

| 

7* 

24 

34 

43 

61 

4 

5 

December ... 

73 

21 

2 9 

49 

67 

3 

4 













244 

not enter now—deficient agglutinin formation, excess of 
agglutinoids, etc. 

^2 That a positive result is in many cases, one might almost 
say in most, not obtained until the end of the fitst week 
of the illness, or even later. 

(3) That in cases of exceptional severity the test may give a 
negative result. 

(4) That a positive result does not necessarily mean that the 
present illness of the patient is enteric fever. Patients 
who have been through an attack previously, it may be 
years before, not uncommonly give agglutination, 
especially if they are carriers, but by no means always so. 

Blood culture, of course, is best for early diagnosis. The 
percentage of cases from which the organism can be isolated 
diminishes from 96 in the first week to 35 in the fourth; that is, 
the value of the Widal test gains as that of the blood cultivation 
diminishes. 

To state briefly the effects of the Widal results on the notification 
returns (in addition to what was stated before): negative results, 
owing to early stage of disease, delayed reaction, and so forth, 
would tend to diminish the number notified, while positive results 
occurring in old, cured cases, suffering now from totally different 
affection, would tend unduly to swell the returns. 

Bearing these few facts and reservations in mind, the charts 
giving the curves of notifications to the Medical Officer of Health 
for this disease for the last seven years are most instructive (see 
Charts II—X, pp. 279-284). 

In nearly all of these charts it will be noticed that there is a 
distinct rise in the number of cases notified in the second quarter of 
the year, and in some years a small increase during November. In 
1912 the November increase was markedly high (see Chart VI), the 
reason for which I am unable to conjecture at this interval of time. 
Comparing the average table of notifications (Chart X) or the 1914 
one (Chart VIII) with the laboratory results for the same year 
(Table I), the first-named increase is most distinct. 

The epidemiological aspect of the question is a fascinating one, 
but appertains rather to the domain of the Medical Officer of 



Health, and I must pass on to the points which belong more to my 
own investigations. 

Chart X reveals the startling fact that for the past seven years 
—I have not been able to obtain reliable statistics prior to that— 
the average number of fresh notifications of enteric fever cases has 
never fallen as low as 15 in any month of the year, while in May and 
June the average for the same period reaches the high figure of 30 
and over. 

Briefly stated, then, the problem to be solved is two-fold, 
namely: — 

(1) Why is the permanent enteric fever prevalence so high in 
a town favourably circumstanced as regards sunlight and 
natural drainage? 

(2) Why is the prevalence at its highest in the late spring and 
early summer months of the year ? 

I am strongly of opinion that the solution of the problem 
of enteric fever prevalence, and thereby the hastening of the 
desideratum of the final suppression of the disease, is more likely to 
be reached by studies of the affection in areas and places where it is 
endemic than by investigations of local acute outbreaks where of 
necessity the investigation has to be, to a certain extent at least, a 
hurried one. A town in which the average notification total in the 
best months is between fifteen and twenty cannot be regarded 
otherwise than as an endemic area. 

In beginning an investigation into the causes of such prevalence 
of enteric fever in Kingston, one’s thoughts were naturally first 
directed to the usually recognised sources of the spread of the 
disease, namely, food and water, flies, and dust. 

II. WATER SUPPLIES 

Bacterial examinations of the Kingston water supplies are 
regularly carried out at this laboratory in order to safeguard their 
purity. The supply is three-fold; of these, two are in constant use, 
while the third is a subsidiary supply called into requisition in dry 
months, when the others prove inadequate for the demands of the 
population. If the analyses at any time give indications of 
deterioration from the usual standard, which is a good one and well 



246 


up to that recognised for tropical waters, that is, if any indications 
of excretal contamination are found, the fact is immediately reported 
to the Central Board of Health and to the Kingston General 
Commissioners who control the supplies, and steps are at once taken 
to locate and remedy the mischief. 

It came as a matter of great surprise, therefore, in June, 1914, 
when the town was scared by a report from one in authority in 
public health matters here, that the water supply, and especially the 
auxiliary one installed comparatively recently at great expense and 
after several careful analyses, consisted of ‘ diluted sewage/ and 
could ‘by no methods be rendered fit for drinking/ and was the 
cause of the large incidence of enteric fever. 

Fortunately, to put the matter briefly, the report proved to be 
quite unfounded, a mere canard, for the incriminated source of 
supply was not, nor had it been for some time, in use, the other two 
sources proving quite sufficient for the demands of the population. 
Moreover, these latter showed no deviation from their customary 
standards. 

To discuss in detail the points brought forward would unduly 
prolong this paper, so I will content myself with stating that against 
the supposition of the disease in Kingston being water-borne are the 
following facts: 

1. The population of Kingston is estimated at 58,352, and the 
notifications of cases of enteric fever include all those in the Public 
General Hospital, which draws not only from Kingston, but also 
from the neighbouring parish of St. Andrew, at all events the lowest 
and most densely populated part of it. At the very smallest 
computation, therefore, we may take it that the combined population 
from which the notifications came was 60,000. 

During May, the month in which the greatest number of cases 
was notified, there were fifty notifications, that is, less than 1 per 
1,000. This is a large proportion in a town with every natural 
advantage for drainage, but a very small proportion as compared 
with what would occur if the infection were water-borne, since over 
58,000 individuals drink this water. The records of the Worthing 
outbreak in 1893 give 557 cases in three weeks in a population of 
15,000, or approximately 37 per 1,000. In the Maidstone epidemic 
of 1897 there were 1,201 cases in three weeks out of a population of 



247 


33.830. or 35 cases per 1,000. In Lincoln in 1905 in the same period 
approximately 11 per 1,000 of the population were attacked. In 
Kingston, as already stated, it is less than 1 per 1,000. 

2. The penitentiary, with a large number of inmates, and the 
lunatic asylum of 1,400 patients, use the same water, and moreover, 
water from no other source, yet amongst these during the same 
period there were no notifications at all. 

3. The different quarters of the town, though having the same 
water supply, contributed in very different proportions to‘the total 
of cases of enteric fever notified. Thus, in the month referred to, 
from the N.E., N.W., S.W. and S.E. districts there were notified 
respectively 11, 23, 7, and 5 cases, that is, 34 in the northern to 12 
in the southern parts. 

In subsequent months for which I have, by the kindness of 
Dr. Cross well, the Acting Medical Officer of Health, been able to 
obtain details, in July, out of a total of 19 for all four districts, 15 
came from the northern and 4 from the southern; in August, 19 out 
of 28 were notified from the same parts; in September, 12 out of 20; 
in October, 11 out of a total of 17; in November, 12 out of 17. The 
significance of these facts will appear later. 

III. MILK AND OTHER FORMS OF FOOD 

Personally, I have not observed any cases in Kingston in which 
the infection could be definitely traced to contaminated milk supply. 
This is a matter for the Medical Officer of Health, and is outside 
my particular province. I am on sure ground, however, in stating 
that here, as in most tropical countries, milk is very carelessly 
handled, and the cleanliness of the vendors and their receptacles is 
by no means above suspicion. Watering the milk is a common 
practice, and if the tap is too far off, a nearer source is called into 
requisition without compunction. One vendor has been seen to 
abstract milk and fill up to the original bulk with liquid flowing 
down the gutter at the side of the road. When it is remembered that 
these same gutters constitute for the poorer inhabitants the State- 
provided latrine, one source of infection is not far to seek. 

Besides the faulty methods of handling and delivering milk, 
another and more remote source of infection is possible. Few, if 



248 


any, of'the purveyors—I personally know of none, except possibly 
at the Government farm—make even a pretence of sterilising milk 
bottles or cans. So infection of milk may arise by washing the 
various utensils with polluted water, by employment of carriers, or 
of persons suffering from mild or ambulant forms of the disease. 

As regards other articles of food, shell-fish is not eaten to any 
extent in Kingston, and certainly not by the poorer classes. 
Vegetables are brought into the town from neighbouring districts by 
small cultivators, who deposit their excreta close to the huts in which 
they live; they form a possible but not a proved source. 

Sweets, cakes, and such like, are on sale at various dusty street 
comers, and it is only recently that the vendors have been ordered 
to keep these delicacies ( ?) under cover to protect them from dust, 
flies, or the fingers of intending purchasers. These people commonly 
take up one article after another before coming to a decision as to 
which gives most value for their money. The hovels in which sweets 
and foodstuffs are prepared are anything but above suspicion. 
Indeed, I know of one case where the sweets thus sold were actually 
being made in a hut in which an enteric patient was lying. This was 
only discovered by the merest chance, and the tracing of infection to 
its source when spread from unrecognised or unreported cases must 
of necessity often be a matter of chance. Nevertheless, it is clear 
that from the standpoint of preventive medicine, the unrecognised 
cases are of greater importance than cases which are erroneously 
reported as enteric fever. 


IV. FLIES 

These insects are very troublesome in some parts of the city, 
particularly in the poorer parts whence many of the enteric cases are 
notified. It is well known, in fact almost obvious, that the chances 
of dissemination of infection by the agency of flies in a well-sewered 
city are much less than in cases where proper care is not taken in the 
disposal of excreta. 

In Kingston the parts unconnected with the water-carriage 
sewage system are those in which the majority of cases arise. In 
these quarters flies are troublesome at most seasons of the year, but 
the time when they become a positive pest is that of the 4 Mango 



249 


season/ starting about May, and this is the time at which the 
enteric fever notifications begin especially to increase. This part of 
the question is barely touched upon here; I hope to consider it 
subsequently. 

A fly census in different parts of the town, and the establishment 
of a correlation between this and the districts whence notifications of 
enteric fever are sent, would be an interesting matter, but is more 
within the province of the Medical Officer of Health than that of 
myself. With his consent, I would like to undertake such an 
investigation at some future period, if my other duties will permit. 

Dust, though often very troublesome in Kingston, is not regarded 
as so potent a cause of the spread of enteric fever as has been 
believed, especially in a tropical country with prolonged exposure 
to the sun’s rays. In fact, it would be more correct to say that as a 
direct cause dust accounts for very few cases, and that many of those 
formerly attributed to dust, as air-borne, may be more reasonably 
ascribed to the agency of flies. 

House-flies have been captured in dwellings near badly-kept 
privies used by enteric fever patients, and the bacilli have been 
isolated from them. Experimental investigations (Hamilton, 1903) 
have shown that living bacilli may remain in or on the bodies of flies 
for twenty-three days after infection. 

Cockroaches also, which, like the poor, are always with us in 
Jamaica, swarm near badly protected food, and may easily act as 
mechanical carriers of infection. Food protection, except from the 
point of view of larceny, does not enter into the domestic arrange¬ 
ments of the poorer inhabitants of the city of Kingston. 


V. SEWAGE DISPOSAL 

A few words will suffice to sum up this section. The water- 
carriage system is laid down for the lower part of the town only. 
The upper parts, N.E. and N.W. districts, are largely furnished— 
more or less badly—with privy middens, dry earth closets, and so 
forth, nothing less than a standing and open invitation to flies, 
which freely avail themselves of the opportunity. 

Some of the huts inhabited by natives in Kingston have the 
luxury of a privy to themselves. In other instances there is one to a 



250 


yard in which there may be four or more huts and as many families, 
living as one small related community—very much related—so that 
privacy is more honoured in the breach than in the observance. The 
gutter passing the front doors is so handy that it is by no means an 
uncommon thing to see children, and even adults, obey the calls of 
nature in the open street. Again, natives come down from the inland 
districts, walk 15 miles or more to bring produce to sell in the town, 
stay the night in Kingston or on the borders of the city, and sleep 
on the pavement at the side of the road. These people cannot be 
blamed for depositing their excreta in the road or in the gutter, for 
public latrines are not provided for them. 

These itinerant vendors may account for a few cases, but if so the 
association would be practically impossible to trace. I think that in 
Kingston, considering the class of patients dealt with in hospital, 
with whom my special investigations shortly to be related were made, 
and the quarters of the town from which they came, there is but little 
importation of the disease from outside. It must arise, therefore, to 
a large extent, if not entirely, in factors whose operation is 
localised. 

It has been shown from observations (by Rosenau, Lumsden, 
Kastle, and Anderson, 1909-11) extending over several years in the 
district of Columbia, U.S.A., that there is in general ‘no striking 
difference in the prevalence of typhoid fever in the sewered and 
non-sewered districts/ but at certain seasons, particularly the time 
when flies are numerous, the non-sewered districts showed a much 
greater incidence. 

Also, the chances of infection being conveyed to the healthy from 
close association with the sick are found to be considerably greater 
amongst those who dwell in residences provided with privies, than 
amongst those whose residences have a water-carriage system 
connected with the regular sewage system of the city. 

In Kingston many of the privies are in a parlous state and far 
below the modem standard of a sanitary privy, if such there be. 
They are not water-tight, they are not protected from flies, even in 
the better houses they are often situated close to the kitchens and 
even dining and sleeping rooms. The kitchens are not screened, 
and flies can travel readily from the privy to the food. Also, as 
already stated, in the poorer quarters one privy may have to serve 



for several families, so that many individuals may become infected 
from a single carrier or a common source. 

There is always a danger, in investigating for one single cause, 
of ignoring or forgetting other contributing causes. Thus, an investi¬ 
gation at one time might elicit evidence pointing strongly to one 
factor, e.g., flies, while at another the evidence might be to the 
incrimination of some article of diet—milk, shell-fish, etc.—and tend 
to minimise the importance of the former. But a high general 
typhoid rate with increased prevalence at some particular season of 
the year makes one think firstly what general and continuously- 
acting factor is at work; and secondly, what factor or factors are 
more strongly operative at the time of greater prevalence, and 
whether the reason is to be discovered in an increased scattering of 
seed, increased receptivity of soil, or both combined. 

In Kingston, where the variations of temperature are compara¬ 
tively small in the different seasons and where perpetual summer 
reigns, the following factors must be considered: prevalence of 
flies at certain times, diminution of rainfall and concentration of 
pollution, or sudden heavy rains washing pollution from the banks of 
a stream into the supply, the ingestion of raw fruits, and the 
prevalence of other intestinal affections rendering the subjects more 
liable to contract typhoid fever when exposed to infection, and so 
forth. 

The water supply, the fly nuisance, the sewage arrangements, 
are matters for the Medical Officer of Health to consider and deal 
with. I could not interfere with them without his full consent and 
co-operation. But even granting that these were unsatisfactory, 
there remains still unaccounted for the source whence the flies, or the 
food, or the dust obtained the organisms of the disease, and carriers 
are naturally suspected. 


VI. CARRIERS 

The question of carriers is a matter with which my work as 
Government Bacteriologist is closely connected, but there were 
certain difficulties in the way of investigation to avoid interference 
with the work of the Medical Officer of Health. 

There are difficulties also as to obtaining material, for persons in 



health cannot be prevailed upon to send excreta in order that it may 
be determined whether or no they are typhoid carriers. The 
objection is quite a natural one, and more particularly would this 
be the case when the finding would lead to certain rules and restric¬ 
tions being imposed. Moreover, it is fairly certain that, whatever 
rules were given and however mild the restrictions, they would be 
evaded in some way or other. A rule without an enforceable penalty 
for disobedience is better left unmade. The native who would be 
found to be so altruistic as voluntarily to place restrictions upon 
himself for the good of others would indeed be a ‘rara avis.’ 
Further, if the results of examination were negative, the case would 
by no means be proven, as the carrying might be intermittent. 

The question of the percentage of enteric fever patients who 
become carriers has been studied by many investigators. The 
results, so far as the literature at my disposal goes, may be briefly 
given. Lentz (1905) states that about 4 per cent, of typhoid patients 
discharge the bacteria in their excreta at least ten weeks after 
convalescence. Schneider (quoted by Besanqon) estimates the 
chronic carriers as 3 per cent, of patients. Lehmann and Neumann 
give 2 per cent, to 4 per cent. Nieter and Liefmann (1906-7) isolated 
7 out of 250 patients (that is, 3 per cent.) in an asylum. Asylums 
generally appear to harbour an abnormally high proportion of 
carriers. 

But cases excreting only for so short a time as ten weeks after 
the illness cannot be regarded as carriers in the real sense of the 
term. The percentage of chronic carriers is variously estimated at 
from i‘5 per cent, to 5 2 per cent. So far as I have been able to 
ascertain, the following figures are authentic: — 

Out of 400 cases of enteric fever 6 excreted the bacilli for more 
than three months, giving a percentage of 15; of 316 cases 9 carriers, 
or 2 8 per cent., were found; of nearly 7,000 cases 166, or 24 per 
cent; while of the large number of 11,000 cases 411 carriers were 
found, a percentage of 37. 

Altogether I have found results recorded of 18,431 patients, and 
of this large number 59 2 were detected as carriers, a proportion of 
3 21 per cent. We shall be safe in taking it that, as an average, 
some 3 per cent, of patients become carriers for three months at 
least. 



253 


Of i,800 healthy subjects reported upon by Kayser in 1907 giving 
no history of previous typhoid fever, 27, or 15 per cent., were found 
to be excreting the bacilli. In Columbia the excreta from 1,000 
healthy persons were examined, and of these three were found to 
contain the organisms; one of these excreted them in the urine, the 
other two in the faeces; all three were males. 

Returning to the condition of things in Kingston, my suspicions 
were that, as the enteric rate is so high, there might be individuals 
going about apparently in good health, who were unwittingly 
spreading the infection, or that the percentage of carriers here is 
greater than has been found to prevail in most places. 

VII. BACTERIOLOGICAL INVESTIGATIONS 

It is now an established fact that during an attack of enteric 
fever the bile frequently becomes infected; this is corroborated by my 
results as shown in the table of cases so examined (see Table II). 

If now, as is also known to occur, an inflammatory process arises 
in the gall-bladder, the bacilli may be discharged either continuously 
or at intervals from the bladder into the intestines for periods of 
almost indefinite duration. 

Progressing a step further, it may be said that in many cases 
the chronic bacillus-carriers are individuals harbouring the typhoid 
bacilli in their gall-bladders. 

I determined, therefore, to make cultivations from the bile of 
every patient dying in the general hospital—regardless of the actual 
cause of death as diagnosed clinically or discovered post-mortem— 
in order to find out whether any, and if so how many, were carrying 
the organisms in their gall-bladders at the time of death. 

Of course, in a large percentage of patients admitted with a 
history of enteric fever and dying from this disease, and also in 
patients wrongly diagnosed, but actually suffering from enteric fever, 
one would naturally expect to find the organism. But there was 
yet another class for which I particularly wished to look out, namely, 
patients who were admitted into hospital for some affection totally 
unconnected, or not suspected of being connected with enteric fever, 
and who gave no history of any previous attack of the disease or of 
any prolonged fever which might be so explained. These persons, 



* 5 + 


by harbouring the Bacillus typhosus in their gall-bladders, had 
possibly, if not probably, been acting as carriers, and, had they 
chanced to recover, would have again gone about spreading the 
organisms broadcast. These were the individuals who constituted 
a danger to the community, and if such were found to exist, the fact 
would go far to account for the undue prevalence of enteric fever in 
Kingston. 

It is but right to call attention to the fact that the mere denial 
on the part of the patient as to his having suffered previously from 
the disease does not exclude the possibility of his having passed 
through an attack. The native is so accustomed to short attacks of 
malarial fever that any ‘ fever * of a prolonged or severe character 
usually makes a permanent impression on his mind. When he speaks 
of a 1 bad fever * lasting for two or three weeks, this is quite sufficient 
to enable it to be stated that the history evinces a strong probability 
of the patient having had an attack of enteric fever in the past. 

The investigation into the results of bile cultivations is still being 
continued. Up to the present 120 cases have been dealt with, 
sufficient for a preliminary note, and sufficient to form a basis for 
discussion (see also Addendum, p. 269). Before entering more 
fully into a consideration of the various points which present 
themselves, I would beg to call attention to the appended list of 
cases where the results are briefly recorded (Table II, pp. 272-278). 

The following points worthy of more detailed mention present 
themselves on a persual of this table: 

1. The number of cases exhibiting post-mortem the signs of 
enteric fever from which a positive result was obtained 
from cultivation of the bile, and arising out of this, the 
number of cases with similar signs from which the result 
was negative. 

2. A corollary to the last, namely, a consideration of the 
cases which showed no signs of enteric fever, but from 
which a positive culture was obtained. 

Other points of interest which declared themselves during the 
investigation were: — 

1. Peculiarities of the isolated organism in certain instances. 

2. Absence in most cases of macroscopic signs of inflamma¬ 
tion of the gall-bladder. 



255 

(i) The method of procedure employed in each case 

As soon after death as possible, before starting on the usual 
post-mortem routine, the gall-bladder was exposed, the surface was 
sterilised by searing, and the organ held by sterile forceps. A small 
incision was made with sterilised scissors. Tubes of peptone broth 
were ready containing quantities of a i in 10,000 solution of brilliant 
green, varying from 003 c.c. to 3 c.c., namely, 0*03, 006, 012, 
0*18, 2, and 3 c.c., the brilliant green being added immediately 
before the tubes were taken to the post-mortem room. A loopful 
(4 mm. in diameter) of the bile was introduced into each tube. Six 
tubes were sometimes used, but on no occasion less than three, 
because it was early noticed that the brilliant green had a variable 
inhibiting power, not always proportionate to the amount added. 
In other words, some strains of the bacillus obtained showed less 
resistance to the brilliant green than did others. 

The tubes after inoculation were placed in the incubator at 37°C. 
and examined on the following day. If there was but a slight 
growth with faint cloudiness of the medium, further examination 
was postponed for another twenty-four hours. If there was no 
growth by that time the tubes were kept for a week before the bile 
was noted down as being ‘ sterile.* 

If, and when, growth occurred a loopful was plated on to 
Rebipelagar for isolation, and suspicious colonies, non-lactose 
fermenters, were transferred to nutrient broth. Secondary tests were 
then carried out with various sugars, namely, glucose, saccharose, 
lactose, maltose, mannite, dulcite, and sorbite. 

Any organism isolated, which gave the usual typhosus reactions, 
was then tested for its agglutination power with an immune serum 
which I prepared, and which gave an agglutination titre of 1 in 
4,000 with its own producing strain. Only those organisms which 
gave a positive result with the serum diluted to about this degree 
were regarded as true Bacillus typhosus. As is well known, to test 
with the serum of a typhoid patient or convalescent is quite 
inadmissible for the identification of a suspected bacillus. In such 
a serum besides the main specific agglutinins for the Bacillus 
typhosus , there may be present a whole series of group-agglutinins, 
agglutinoids and iso-agglutinins, in such concentration a$ to con¬ 
stitute a source of gross errors. 



2 5 6 


In the investigation, control tests were carried out with 
corresponding dilutions of normal serum of the derivative animal, 
to exclude the chance that the agglutination observed with the 
immune serum might be an action of a normal serum on the strain of 
bacterium under examination. 

A second control was also put up, using normal saline in place 
of the serum,' in order to exclude any incidental agglutinating 
action of the diluent—a pseudo-agglutination—which might appear 
from spontaneous clumping or insufficient emulsification. 

Bruns and Kayser (1903) propose to use only a low dilution, 
such as 1 in 100 of the immune serum, but this course is liable to 
lead to erroneous conclusions, which need not be fully discussed 
here. A long series of dilutions has been made in each of my cases 
in order to determine in what degree of dilution of the serum the 
agglutination would take place. As Pfeiffer, Lipschiitz (1904), 
and others have pointed out, only when this corresponds fairly 
approximately with the estimated titre of the serum ought the 
a ggl u ti na tio n to be regarded as evincing the true typhosus nature 
of the organism in question. 

(ii) Cases with post-mortem signs of enteric fever from which a 
positive result was obtained on cultivation of the bile. 

Out of the one hundred and twenty cases, twenty-eight were 
shown at the autopsy to be suffering from enteric fever, that is to say, 
definite macroscopic lesions of the disease were present. In twenty- 
four of these the Bacillus typhosus was isolated from the bile, and 
in one other, whose blood during life had given a positive agglutina¬ 
tion with B. paratyphosus A, and negative with B. typhosus , and 
who presented symptoms typical of enteric fever, the B. para¬ 
typhosus A was isolated. We may therefore say that from twenty- 
eight cases of enteric fever the causative organism was isolated from 
the bile in twenty-five, or in 89 28 per cent., although I am aware 
that the reckoning as a percentage with so few cases as twenty-eight 
is liable to error. This percentage number agrees almost exactly 
with the figures of Forster and Kayser (quoted by Hewlett), who 
obtained pure cultures from the gall-bladders of seven out of eight 
cases. A better comparison may be made between these cases and 



257 


mine if we divide my twenty-eight into series of eight. For the 
first eight I obtained the B. typhosus in pure culture six times, and 
twice the organism was associated with a lactose-fermenting one, 
which proved to be the ordinary Bacillus coli. The B. typhosus 
was thus isolated in all the first eight. One of the cases in which 
the B. coli was present also died from general peritonitis following 
perforation of a typhoid ulcer, and the bile may have become 
infected by general systemic infection, or, what is an equally 
probable explanation, the B. coli may have entered as a contamina¬ 
tion in making the culture. If the former explanation is the correct 
one, the fact shows that the bile and brilliant green do not always 
have an inhibiting action on Coli organisms. 

With regard to the second series of eight, I obtained the 
B. typhosus in pure culture in six, and B. paratyphosus A in 
another, also in pure culture; in other words, the organisms were 
present in pure culture in seven out of eight. In the third series, 
B. typhosus was obtained in pure culture in six, one remained 
sterile, while from the other B. coli only was obtained. 

The remaining four of the twenty-eight yielded the B. typhosus 
alone in each case. 

Hiss and Zinsser (1910) in their text-book merely state that the 
‘ typhoid bacilli have been frequently observed in the gall-bladder at 
the autopsy. 1 They do not mention how frequently. According to 
Meyerstein (1907) the bile of typhoid patients nearly always 
contains the Bacillus typhosus , though its presence does not often 
give rise to clinical symptoms. Its presence may, however, set up 
inflammatory processes in the wall of the gall-bladder. Thus 
Stewart (1901) found a condition of cholecystitis in about 1 per 
cent, of cases. Similar findings have been reported by Forster and 
Kayser (1905), Hilgermann (1909), Findlay and Buchanan (1906), 
Marmann (1908), Nieter (1906-7), and others. 

The presence of the bacilli in the gall-bladder of patients 
suffering from enteric fever is not to be wondered at when we 
consider that the disease is, in the early stages at least, a 
bacteriaemia; that experiment has shown that it is by way of the 
blood-stream that the gall-bladder becomes infected, and that the 
conditions prevailing there appear to have a favourable influence 
upon the growth of this organism. For it has been shown by 



Blackstein and Welch (1891) that after intravenous injection of the 
bacilli into guinea-pigs they were recognisable in the bile four 
months later, when all other organs appeared to be completely free 
from them, while after subcutaneous, intraperitoneal, and intra- 
gastric inoculation the bile remained sterile, though it was infected 
after intravenous injection, even with the cystic duct ligatured 
(Doerr, 1905). This appears to be the usual mode of infection of 
the bile in man, namely, by way of the circulation. It is only fair, 
however, to mention that Koch and Chiarolanza (1909) have held 
that there is an alternative route direct from the intestine to the bile 
through the ductus choledochus, at all events in experimental 
infection of rabbits. In the case of the human subject, however, 
this latter is very unlikely, seeing that the lower part of the small 
intestine is that most affected, and it would indeed, with respect to 
the local condition, be a 4 long way to go* when the route via the 
blood-stream is so much more accessible and rapid. Moreover, the 
bile appears to be infected so very early in the disease. 

(iii) Cases which showed no post-mortem evidence of enteric fever , 
and in which no history of such was obtained , but which t 
nevertheless , yielded a positive result on cultivation of the 
bile. 

This group is of the greatest importance in lending support to 
the suspicion on which the investigation was undertaken, namely, 
that unrecognised possible carriers are going about in Kingston in 
larger proportions than have been estimated in other countries. 

As has been already stated, some 3 per cent, of patients 
become carriers for a considerable time. Of this first series of 
one hundred and twenty autopsies there have been found four who 
up to the time of onset of their final illness had been going about 
apparently in perfect health, who gave no history of having had an 
attack of typhoid fever. In one case, No. 2, there was an indefinite 
history of prolonged fever which might possibly have been typhoid, 
though from the symptoms and post-mortem signs this might equally 
well be ascribed to tuberculosis. The four mixed freely with their 
fellows, and lived in the poorer, insanitary, and unsewered parts 
of the city. 



*59 


These cases are worth quoting briefly: 

1. C. C., male, aged 49 years, white, No. 7 in the appended 
list, Table II. Admitted to hospital 4 p.m. March 31, 1914* 

a history of having at 9 a.m. ‘taken rat-poison in mistake for 
phenacetin.' He died atr 11.40 p.m. The fact that more than 400 
grains of arsenious acid were found in the stomach and viscera, even 
after the vomiting which had continued for ten hours, points to 
deliberate suicide, for no one would take nearly an ounce ol 
phenacetin for a single dose. His neighbours stated that he went 
in and out amongst them apparently in perfect health till the day 
of his death. In bile culture B. typhosus was obtained. 

2. B., a Coolie woman, aged 21 years, No. 34 in the list, when 
admitted complaining of ‘fever, cough, and pain in the right side 
of the chest.' Expiration was prolonged, and there were rhonchi 
audible over both lungs. 

The diagnosis of phthisis was made, and the patient died early 
the following morning. 

At the autopsy the lungs showed minute scattered nodules with 
caseous contents, possibly tuberculous, but the chief finding of 
interest was that the gall-bladder had practically disappeared, and 
there was an ulceration into the duodenum just beyond the pylorus, 
with a gall-stone about the size of a small hazel nut in the aperture. 

There were three abscesses in the liver, one cavity being the size 
of a tangerine orange. 

From the nucleus of the gall-stone, the only one found, a pure 
culture of B. typhosus was obtained. 

3. S. D., male, aged 19 years, black, No. 73 in the list, was 
admitted with a history of six days* ‘ cough, pain in the right side 
of the chest, and fever.' The percussion note over the right lung, 
lower lobe, was quite dull. ‘ No breath sounds could be heard, but 
vocal fremitus and resonance increased.' Widal reaction was 
negative. The patient died ten days later. 

At the post-mortem the right lung was quite solid; the upper 
and middle lobes in a state of grey hepatisation, the lower lobe in 
a condition of purulent infiltration, and in one place broken down 
to abscess formation. The lung was large and heavy, and the liver 
so displaced that the upper margin only reached one and a half 
fingers'-breadth above the costal edge. 



2 6 o 


This, then, was a case of unresolved lobar pneumonia. No 
history whatever was obtained pointing to enteric fever, but a culture 
of B. typhosus was yielded by the bile. 

4. E. R., female, aged 21 years, black, No. 113 in list, was 
admitted to hospital on October 26th, 1914, with dyspnoea fend 
general oedema, especially of the lower limbs. The urine was 
albuminous and contained granular and hyaline casts. Death took 
place on the following day. 

The autopsy revealed a condition of chronic nephritis, the renal 
capsules were adherent, the surface of the organs granular, the 
cortex narrowed to about half the normal ratio, and there were two 
small cysts in it. The heart was enlarged, the left ventricle being 
hypertrophied, and the weight of this organ when empty was 480 
grams. There was no sign of enteric fever found, and no 
history obtained of any such disease. A pure culture of B. typhosus 
was isolated from the bile, which agglutinated ‘after three sub¬ 
cultures with the typhoid immune serum in as high a dilution as 
1 in 5,000. 

Exclusive, therefore, of cases showing evidence of enteric fever 
at the autopsy, the bacillus has been isolated from the bile of four 
subjects: 

1. Dying from arsenic poisoning, suicidal. 

2. Dying from multiple abscesses of the liver. 

3. Dying from pneumonia. 

4. Dying from chronic nephritis and heart disease. 

It would hardly be fair to draw conclusions from so few cases as 
one hundred and twenty, and the investigation is being continued 
(see Addendum, p. 269). All that can be said at present is that, 
apart from patients treated for enteric fever at the hospital, and 
apart from cases showing signs of this disease post-mortem, there 
have been three among the first one hundred and twenty who were 
harbouring the Bacillus typhosus in their gall-bladders at the time 
of death. In a fourth who had done so, the organism was found in 
a gall-stone which had ulcerated into the duodenum. 

If we deduct the number of those who were suffering from the 
disease, showing definite signs of it at the autopsy, we may state 
that out of ninety-two subjects there were four who were harbouring 
the organism of enteric fever. 



26i 


(iv) The deductions which may be drawn from the tact of isolation 
of the bacillus from the gall-bladder or its contents 

1. The subjects are chronic typhoid carriers. This is possible, 
but could not be regarded as proved unless repeated examinations 
of the excreta yielded positive results. In my cases this was 
impossible, as they did not come under my observation till after 
death. 

2. The subjects are temporary carriers. 

(i) They may, in the absence of history, have passed through 
an attack of enteric fever and forgotten or not have known the fact, 
either because of the mildness of the infection, or because of time 
elapsing, or because the illness had been wrongly diagnosed. The 
following cases show that the last may certainly occur: 

No. 4, diagnosed lobar pneumonia. 

No. 5, diagnosed pneumonia. 

No. 9, diagnosed pneumonia. 

In these three, typhoid ulceration was found but no signs 
of pneumonia. 

No. io, diagnosed general peritonitis. This was true, but 
the peritonitis arose from the perforation of an enteric ulcer 
which had not been suspected. 

No. 88, diagnosed cerebral haemorrhage. There were the 
usual signs of enteric fever, but none whatever of cerebral 
haemorrhage. 

No. 99, diagnosed pneumonia. This was present, but there 
had been no suspicion of enteric fever, which was evident with 
its typical pathological changes. 

(ii) They may be ‘ porteurs sains * in the sense of never having 
had the disease. In this connection I may quote an interesting case 
which came to my notice as Government Bacteriologist more than 
a year ago. It was mentioned in my paper already referred to, and 
published in the ‘Practitioner* for November, 1913: 

‘ A child of nine months suffered from an attack of typhoid fever, proved by 
isolation of the bacillus. The question then arose as to the method by which 
the child could have become infected. The little patient’s mother was the nurse 
in charge of the enteric ward at a general hospital in the island, and it was thought 
that she might possibly have taken some milk or other food from the ward home 
to the child. A specimen of the mother’s blood was asked for. and it was found 
to give a very marked agglutination of bacillus typhosus in high dilution. 



262 


‘ The nurse was perfectly certain that she had never suffered from typhoid 
fever, or, in fact, from a prolonged fever of any kind ; she had always been healthy, 
and “ never remembered being ill in her life ” (to quote her own words). The 
next step was the examination of her stools and urine, and the bacillus typhosus 
was isolated from the latter ; in short, this nurse was definitely a carrier, though 
never having herself suffered from the disease, and she seems without doubt to 
have conveyed the disease to her child.’ 

If the subjects are ‘porteurs sains * in the sense mentioned above, 
that is, the organisms are present without setting up the disease, we 
may find here support for the X Y Z theory of Pettenkofer. Thus, 
it is possible that the bacilli may be harboured in certain situations 
(in my cases the gall-bladder) for a lengthy period, but until the 
other components of the aetiological complex are superadded no 
disease results. We may, perhaps, compare it to the lowering of 
resistance to the Bacillus tuberculosis , allowing development of the 
organism and production of the disease. 

Against this in my cases is the fact that two, possibly three, of 
the subjects were considerably debilitated by illness, and there were, 
nevertheless, no signs of enteric fever supervening. This argument, 
it must be confessed, is of the nature of a two-edged sword, for, 
firstly, mere debility by any illness may not be the Y or Z of the 
aetiological complex, but some element more specific in nature may 
be necessary. Secondly, some of the cases which showed signs of 
enteric fever post-mortem, although a different diagnosis had been 
made clinically, might be instances of the supervention of enteric 
on a previous debilitating disease. This, again, opens up the 
interesting question as to whether cases of enteric fever arising in 
patients who have been in hospital for some time, a month perhaps, 
may not in some instances be of f spontaneous * or ‘ autogenous * 
origin, instead of being regarded as cases of ‘ contraction of disease 
in hospital/ By the terms spontaneous or autogenous I imply the 
‘ porteurs sains * who develop the disease after some other debilitating 
illness, for example, typhoid fever following dysentery, and not 
merely ordinary cases with a prolonged incubation period. 

(v) Peculiarities of the organism isolated in some instances . 

1. Motility. As has been noted by Fischer (1909), I have found 
several times that when freshly isolated from the body, that is, from 
the bile in my cases, the bacilli were but slightly motile, and in some 



263 


instances were devoid of motility altogether. But he goes on to 
say that even after prolonged subculture the motility was only 
slight. This has not been my experience. After the third or fourth 
transference to nutrient broth the normal motility was always 
regained; or by transference from agar to broth alternately. 

I have not so far had to employ any of the special media for this 
purpose, such as Losener’s serum, Terni’s peptone-free glycerin 
bouillon, or the 2 per cent, glucose broth of Germano and Manrea. 

2. Agglutinability. In my cases I have not yet met with an 
organism which, while giving the cultural tests and sugar reactions 
of the B. typhosus , was completely non-agglutinable with the 
prepared immune serum. In several instances, however, the first 
subculture showed very slight agglutinability, but, after sub¬ 
culture to the third or fourth generation and subsequently, the 
organisms proved normally agglutinable. 

In two cases I found a mixture of two typhosus colonies, one 
easily agglutinable, the other only agglutinating with difficulty, 
occurring from the same subject. A further point was also 
observable, namely, that the former—the one readily agglutinable— 
showed marked motility, while the latter—the poorly agglutinating 
strain—was practically non-motile. Sub-cultivation, however, on 
ordinary agar and in broth alternately, produced organisms equally 
motile and equally agglutinable in the two cases. Thus, it is seen 
that the diminution or absence of motility and a low degree of 
agglutinability of a newly isolated strain of the organisms go hand 
in hand. By further observations it was found in carrying out the 
agglutination reaction that time was an important factor in some 
cases. Thus, a strain which proved to be agglutinated to a slight 
degree only within the ordinary time limit of two hours, would in 
some cases, not in all, be strongly agglutinated if left for several 
hours—up to 24—at room temperature, less if kept in the incubator 
at 37 0 C. 

Two of the explanations, which have been put forward to account 
for the low degree of agglutinability of typhoid bacilli freshly 
isolated from the body, are worthy of being mentioned in greater 
detail; namely, that of Bail (1901), Muller (1903), Kirstein (1904), 
and others, and that of Nicolle and Trenel (1902). 

Their findings, some of which I have been able to confirm, may 
be briefly stated thus : — 



264 

A culture of typhoid bacilli agglutinable in high dilution of the 
immune serum was inoculated intraperitoneally into a guinea-pig. 
After remaining a few hours the agglutination test was again 
applied, and the titre of agglutination was reduced from the previous 
1 in 4,000 almost to nil; there was incomplete clumping even with 
1 in 300. This has been explained by supposing that the bacteria 
which have been inoculated seize upon the agglutinophores which 
possess the haptophore groups but not the zymophore, and thus put 
them out of action. Bail (1901) similarly explains the reduction of 
agglutinability of bacilli growing in a diluted immune serum, for the 
normal degree of agglutinability is restored after a few transferences 
back to ordinary nutrient media. It is but right to state, however, 
that Muller (1903), Hamburger, and others regard the latter 
phenomenon rather as a definite attempt at immunisation of the 
bacillus against the injurious action of the immune serum, though 
they are all agreed as to the subsequent restoration to normal agglu¬ 
tinability. It must be noted that this reduction is not marked in all 
strains. We may thus briefly state this view: Under certain 
conditions typhoid bacilli present in a patient seize upon the 
agglutinoids and so lead to the production of cultures of the 
organism which are but poorly agglutinable. 

The second explanation rests upon the findings of Nicolle and 
Trenel (1902) that cultures of the Bacillus typhosus , normally motile 
and readily agglutinable, lost these properties when grown at 42 0 C., 
though they regained them on subsequent cultivation at 36° C. 
They inferred that the high temperatures which some typhoid 
patients exhibit might produce a similar change in the bacilli, 
reducing their agglutinability. Higher temperatures still, such as 
50°C., on the other hand, appear to bring about a hastening of the 
agglutination and to render it more complete. 

Eisenberg and Volk (1902) have shown by their experiments that 
the non-agglutinable or poorly agglutinable strains may be, 
nevertheless, identified by means of the immune serum. Heating the 
cultures of typhoid bacilli or treating them with weak acids deprived 
them of their agglutinability. Bacilli so treated are capable of 
binding large amounts of agglutinin, hence the inference that such 
treatment renders nugatory that part of the agglutinable substance 
which is actively concerned in agglutination. The haptophore 
group, however, is still present, for it is found that if such altered 



z6s 


bacilli be brought into contact with a typhoid immune serum all the 
typhoid agglutinin can be removed, so that it will no longer 
agglutinate any typhoid bacilli. This, as Wassermann has 
suggested, might be usefully employed for the identification of 
typhoid bacilli in those cases in which there is a suspicion that the 
organisms have, owing to extraneous influences, lost their normal 
agglutinability. But, seeing that cultivation again on ordinary 
nutrient media soon restores the normal agglutinability, this method 
of Wassermann would, or need only, find application where a 
speedy diagnosis is urgently called for. 

(vi) Length of stay of the organism in the gall-bladder. 

As regards the length of time during which the bacillus can exist in 
the gall-bladder, the limit has not yet been fixed. Cases of 16, 17, and 
20 years have been reported by Zinsser (1908), Droba (1899), and 
Hunner-Writer (1905), respectively. I myself related the case of a 
faecal carrier of Bacillus paratyphosus A in 1911, whose primary 
attack occurred in 1889, twenty-two years previously. Cases 
reported by Dupre, Ramond, and Faitout, are quoted by Besan^on, 
but it must be remembered that all these refer to the duration after 
an attack of the disease. The four mentioned in my table, Nos. 7, 
34, 73» and 113, gave no history of any previous attack at all, and 
the time in them is impossible to determine. 

(vii) Presence of the bacillus with and without obvious inflam¬ 
matory changes in the gall-bladder. 

Doerr (1905) has stated that the bacilli make a prolonged stay 
in the gall-bladder only if an inflammatory condition of the mucous 
membrane is set up. In two of my four cases I could not make out 
any such condition. It is possible, therefore, that their histories 
were correct, that they had not suffered from the disease and that 
the gall-bladder was merely their temporary sojourning place, a 
port of call, as it were, in a ‘porteur sain,* as the bacilli were on 
their way to being excreted. It may be incidentally noted that 
gall-stones are very rarely found post-mortem here. Of all the 
autopsies carried out by me during the last four years, I have only 
found them to be present in two instances. 



266 


(viii) The question of atypical cases without characteristic changes. 

Some of those from whom the organism was isolated, although 
no symptoms were present and in whom there were no post-mortem 
sign of the disease, might, of course, be examples of the atypical 
cases without characteristic changes. Under this heading would 
come such as have been reported as cases of 1 typhoid septicaemia * 
If by this term we understand 1 a systemic affection in which the 
causative organism not only gains entrance to the blood-stream but 
also multiplies therein/ I know of no recorded positive findings of 
such a thing in enteric fever, but an ordinary bacteriaemia is the 
normal condition at an early stage of the disease. Absence of 
evidence of intestinal affection, in spite of finding the bacilli in the 
blood, is less uncommon. Such may be explained by the fact that 
all transitions occur between the marked pathological changes in the 
intestines and a minimal, scarcely recognisable lesion, and 
this also apart altogether from the severity of the case. Also 
de Grandmaison (1900) and Krokiewicz (1908) have produced 
evidence that in several such cases definite formation of specific anti¬ 
bodies may be recognised in the serum. A second explanation is 
that mentioned in the last section, namely, that the bacilli have 
gained entrance into the body of a patient suffering from totally 
different disease, the subject acting the part of a passive carrier or 
receiver, as, for example, the entrance into the blood-stream of 
typhoid bacilli accidentally by way of a tuberculous enteritis, as in 
a case recorded by Busse (1908). 

We must never lose sight of the fact that enteric fever is not an 
intestinal infection. It is, on the contrary, a general one with 
localisations of the bacteria in various situations, most frequently in 
the intestine, but often in the liver and biliary system, sometimes in 
the lungs, the bladder, the pleura, the meninges, and so on. Thus, 
there may be a considerable variety ranging between a general 
bacteriaemia without local infection specially marked in the 
intestines, and more definitely localised forms with symptoms of 
meningitis, broncho-pneumonia, cholecystitis, nephritis, arthritis, 
etc. Yet again the typhoid nature of the condition may be 
masked by accompanying infection, such as a streptococcal, or 
tuberculous one. 



VIII. PERSONAL CONTACT 


With the histories often faulty, sometimes unintentionally, some¬ 
times deliberately so, it is an almost impossible matter to trace the 
extent of personal contact and gauge how far it plays a part in 
spreading the disease in Kingston. The poorer inhabitants live 
often closely massed together, and many cases may thus arise from 
a common source of infection, as has been already stated. 

When the large number of possible sources of infection furnished 
by typhoid carriers and by typhoid patients are considered, and 
the many ways in which infection may be conveyed from the diseased 
to the healthy, it can only be wondered why there is not more 
typhoid fever rather than why there is so much. 

If the extensive prevalence of this disease is to be coped with in 
anything like an adequate manner, typhoid convalescents must be 
subjected to greater restrictions than they are at present. 
Particularly is this necessary with cases who pass cloudy urine 
(bacilluria). They must be restricted until their excreta have been 
examined bacteriologically and found to be negative. They must 
not be freed entirely from observation even then, because of the 
possible intermittency of the condition. As everyone knows, it is 
the usual custom to discontinue all supervision over enteric fever 
patients as soon as they are able to leave their beds. Consequently 
in the majority of cases (I know there are exceptions, for instance 
in the army, but otherwise the exceptions are rare) disinfection of 
stools and urine is stopped as soon as the convalescent is able to be 
up and about. To recognise how very unsafe this procedure is, 
we have only to call to mind the results of examination of the excreta 
of convalescents, and note the length of time during which many of 
them continue to harbour the organism in question. 

Though it is of great importance and value to discover the main, 
or even one of the main, factors causing the spread of enteric fever 
in a district, particularly in a crowded town like Kingston, 
nevertheless, since the typhoid rate of incidence may be looked upon 
as a good sanitary index of the hygienic condition of a community, 
coincident improvements should be carried out in all the branches of 
sanitation, if the prevalence is to be materially reduced. The 
purity of the water supplies must be safeguarded, the sewerage 



268 


system should be modernised and the food supplies protected. The 
patients should be kept under observation for a more prolonged 
period than under present circumstances here (and so far as I am 
aware the same remark applies elsewhere). More care should be 
exercised both as to site for and manner of disposal of refuse. 
Measures should be taken to minimise the prevalence of flies. The 
community should be educated to greater personal cleanliness. In 
short, all the conditions of general sanitation should receive attention 
where such are found to be defective. 

IX. LEGISLATIVE RECOMMENDATIONS 

Having pointed out some of the various ways in which the undue 
prevalence of enteric fever may be accounted for in Kingston, it only 
remains for one to suggest means by which the possibility of spread 
may be to a large extent controlled. The following brief recommen¬ 
dations do not include the measures which have been already 
indicated, but apply rather to the legislative aspect of the question. 

1. Rules should be made for early notification of all suspicious 
cases. Much valuable time is lost if the physician waits 
until he is practically certain, often a matter of a week and 
sometimes of a fortnight or more. The form of notification 
should be different from the usual one of 1 In my opinion 
A.B. is suffering from enteric fever/ and might read thus: 

‘ In my opinion A.B. shows evidence of symptoms which 
may indicate enteric fever in an early stage/ Seeing that 
the diagnostic ability of physicians is so variable, and 
that the symptom-complex regarded as pointing to enteric 
fever is so differently interpreted by individual medical 
men, the regulation might be made that any patient with 
unexplained fever of three days’ duration should be so 
reported. 

2. Blood cultures, if the case is in an early stage, or Widal’s 
reaction after, say, the end of the first week should be 
carried out at the Pathological Laboratory free of charge 
to the patient. 

3. Isolation of the patient should be required. In circum¬ 
stances where this cannot be carried out at home to the 



269 


satisfaction of the health officer, removal to hospital with 
this end in view should be made compulsory. 

4. If the patient is allowed to remain at his home, the health 
officer must be satisfied that adequate disinfection of the 
excreta not only can be, but is carried out. 

5. No person continuing to live in the same dwelling as an 
enteric patient should be allowed to prepare, or manufac¬ 
ture foodstuffs (including beverages) without the permission 
of the health officer, given in writing. 

6. Any room which has been occupied by an enteric fever 
patient must be properly disinfected before being again 
occupied. 

7. Patients should remain under observation of the health 
officer until the excreta show on three consecutive examina¬ 
tions, at intervals of a week, absence of the causative 
organism. 

8. All patients on discharge from hospital should be warned of 
the danger they constitute to others. Any infringement of 
the rules, as to abstaining from the preparation, etc., of 
food for others, must be made punishable. 

9. Persons engaged in the manufacture, purveyance, or sale 
of food or drink, must, if called upon to do so by the 
health officer, furnish evidence that he or she is not a 
carrier. For this purpose he or she must allow examina¬ 
tions of the excreta to be made. If the results are negative 
a certificate is to be given to that effect which will hold 
good for a certain time, unless there are definite reasons 
to suspect that the person is spreading infection before the 
time for the next certificate becomes due, when an inter¬ 
mediate examination may be called for. 

10. All such examinations must be carried out free of charge. 


ADDENDUM: Received. 14 April, 1915 

Since the above was written, eighty more cultivations have been 
made; from two of these the Bacillus typhosus was obtained in pure 
culture. Thus, from the total of 200 there have been six cases from 
which the bacillus has been isolated, apart from those subjects who 



270 


showed post-mortem signs of enteric fever. Of these two, one was a 
patient dying of dysentery, the other of tuberculosis of the lungs and 
pleurae. In addition, therefore, to the usually recognised carriers, 
who have definitely passed through an attack of typhoid fever, I 
have found 3 per cent, of persons dying from some other affection 
than typhoid, from whom no history was obtained of having suffered 
from the disease, nevertheless harbouring the bacillus in their gall¬ 
bladders at the time of death. H. H. S. 


REFERENCES 


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Med. Wocbenscbr., Nos. 7. and 12. 

Besan^on, F. (1910). Precis de Microbiologic clinique, p. 256. Masson et Cie, Paris. 

Blachstein, A. G., and Welch (1891). Johns Hopkins Hosp. Bull. II, p. 96. 

Browning, C. H., Gilmour, W., and Mackie, T. J. ( 1913 ). The Isolation of Typhoid bacilli 
from faeces by means of brilliant green in fluid medium. Journ. of Hygiene, XIII, 
PP* 335-34^. 

BrOckner (1910). Ueber Nachuntersuchungen bei Personen, die vor Jahren Typhus durch- 
gemacht haben. Arb. a. d. Kais. Gesundbeitsamt , XXXIII, pp. 436-442. 

Bruns, H., and Kayser, H. (1903). Ueber die Verwertbarkeit des Agglutinationsphanomcns 
zur klinischen Diagnose und zur Identiflzierung von Bakterien der Typhus-Coligruppe 
(Paratyphus u.s.w.). Zeitscbr. /. Hyg., XLIII, pp. 401-425. 

Busse, O. (1908). Ueber das Vorkommen von Typhusbazillen im Blute von nicht Typhus- 
kranke Personen. Munch. Med. Wocbenscbr., LV, pp. 1113-1116. 

Chiarolanza, R. (1909). Experimentelle Untersuchungen iiber die Beziehungen der Typhus¬ 
bazillen zu der Gallenblase und den Gallenwegen. Zeitscbr. f. Hyg., LXII, pp. 11-32. 

De Grandmaison (1900). Arcb. de med. expSr. et <Panat. patbol ., p. 289. 

Doerr, R. (1905). Experimentelle Untersuchungen iiber das Fortwuchem von Typhusbacillen 
in der Gallenblase. Centralbl. f. Bakt., XXXIX, pp. 624-634. 

Droba, S. (1899). Der Zusammenhang zwischen Typhusinfection und Cholelithiasis auf 
grund eines in der Klinik operierten Falles. Wien. Klin. Wocbenscbr ., XII, pp. 1141-1145. 

Dupr£, Ramond and Faitout. Quoted by Bezan^on, Precis de Microbiologie clinique, p. 255. 

Eisenberg, P., and Volk, R. (1902). Untersuchungen iiber die Agglutination. Zeitscbr. 
f. Hyg., XL, pp. 155-195. 

Faitout, F., and Ramond, P. (1896). Angiocholecystite a bacille d'Eberth. Compt. Rend. 
Soc. Biol., pp. 1130-1132. 

Findlay, J. W., and Buchanan, R. M. (1906). Case of Typhoidal Cholecystitis in which the 
usual symptoms of typhoid were absent and in which the bacillus typhosus was isolated 
during life from the cystic and intestinal contents. Glasgow Med. Journ., LXV, 
pp. 177 - 193 * 

Fischer (1909). Klin. Jabrb., XXII. 

Fornet, W. (1912). Statistisches iiber den Typhus und die Typhusbekampfung im Sudwesten 
des Reichs. Arb. a. d. Kais. Gesundbeitsamt, XLI, pp. 448-502. 

Forster, J., and Kayser, H. (1905). Ueber das Vorkommen von Typhusbazillen in der Galle 
von Typhuskranken und 4 Typhusbazillentragem.’ Munch. Med. Wocbenscbr., LII, 
pp. 1474 - 1477 * 



271 


Fxoick (1908). Klin. Jabrb., XIX. 

Hamilton, A. (1903). Journ. American Med. Assoc., XL, p. 576. 

Hewlett, R. T. (1908). Manual of Bacteriology, 3rd Edition, p. 335. 

Hilgermann (1909). Kin. Jabrb., XXI. 

Hiss, P. H., and Zinsser, H. (1910). Textbook of Bacteriology, p. 411. 

Hunner-Writer (1905). Wien. Klin. Wocbenscbr., p. 1476. 

Kayser, H. (1907). Ueber Untersuchungen bei Personen die vor Jahren Typhus durchgemacht 
haben und die Gefsihrlichkeit von 4 Bazillentragem.’ Arb. a. d. Kais. Gesundbeitsamt, 
XXV, pp. 223-228. 

Kirstein, F. (1904). Ueber Beeinflussung der Agglutinerbarkeit von Bakterien, inbesondere 
von Typhusbacillen. Zeitscbr. f. Hyg., XLVI, pp. 229-260. 

Koch, J. (1909). Typhusbazillen und Gallenblase. Zeitscbr. f. Hyg., LXII, pp. 1-10. 

Krokiewicz, A. (1908). Typhusbazillen im Blute und Gruber-Widalsche Reaktion bei Pyamie. 
Wien. Klin. Wocbenscbr., XXI, pp. 1633-1634. 

Lehmann, K. B., and Neumann, R. O. (1912). Lehrbuch der speziellen bakteriologischen 
Diagnostik, Part II, 5th edit., p. 324. 

Lentz (1905)- Klin. Jabrb., XIV, p. 475. 

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Hilfe des v. Drigalsld-Conradischen Nahrbodens und der Agglutination. Centralbl. 
f. Bakt., XXXV, pp. 798-811. 

Lumsden, L. L., and Anderson, J. F. (1911). Report No. 4, on the Origin and Prevalence 
of Typhoid Fever in the District of Columbia (1909-1910). Bull. 78, Public Health 
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Marmann, J. (1908). Bericht iiber die Tatigkeit des bakteriologischen Untersuchungsamts 
zu Gottingen im Jahre 1907-08. Hyg. Rundschau, XVIII, pp. 1013-1020. 

Meyerstein, W. (1907). Ueber die bakteriologische Bedeutung der Gallensalze. Centralbl. 
f. Bakt., XLIV, pp. 434-4+0. 

Muller, P. Th. (1903). Ueber die Immunisierung des Typhus bacillus gegen spezifische 
Agglutinine. Munch. Med. Wocbenscbr ., L, pp. 56-61. 

Nicolle, C., and Trenel, M. (1902). Recherches sur le phfcnomine de Pagglutination. 
Ann. Inst. Pasteur, XVI, pp. 562-586. 

Nieter, A., and Liepmann, H. (1906). Ueber bemerkenswerte Befunde bei Untersuchungen 
auf das Vorhandensein von Typhus bazillentragem in einer Irrenanstalt. Munch. Med. 
Wocbenscbr., LIII, pp. 1611-1612. 

Nieter, A. (1907). Ueber das Vorkommen und die Bedeutung von Typhusbazillentragem in 
Irrenanstalten. Munch. Med. Wocbenscbr., LIV, pp. 1622-1624. 

Pies, W. (1907). Untersuchungen iiber die Wachstumsgeschwindigkeit der Typhusbazillen 
in Galle. Arcbiv. f. Hyg., LXII, pp. 107-127. 

Rosenau, M. J., Lumsden, L. L., and Kastle, J. H. (1909). Report No. 3, on the Origin 
and Prevalence of Typhoid Fever in the District of Columbia (1908). Bull. 52, Public 
Health and Marine Hospital Service of United States, Washington. 

Scott, H. H. (1913). Widal’s Agglutination Reaction as an aid in prognosis in Enteric Fever, 
based on a study of over 1,500 consecutive blood examinations. Practitioner , Nov. 1913, 
pp. 589-608. 

Scott, H. H. (1911-1913). Report of Bacteriological Department, Annual Report Jamaica 
Medical Service, 1911-12. Also Annual Report, 1912-1913. 

Scott, H. H. (1912). Report to the Tropical Research Committee, pp. 30 et seq. 

Stewart, J. (1901). A Report on 620 Cases of Typhoid Fever. Brit. Med. Journ., I, 

pp. 1463-1467. 

Totsuka, K. (1903). Studien iiber Bacterium coli. Zeitscbr . /. Hyg., XLV, pp. 115-124. 

Wassermann, A. von (1902). Ueber Agglutinine und Pracipitine. Zeitscbr. f. Hyg., XLII, 
pp. 267-292. 

Zinsser, H. (1908). A Case of Typhoid bacilli in the Gall Bladder. Proc. of New York Pathol. 
Soc ., pp. 78-81. 



Table II.—Post-mortem findings and results of cultivation of the Bile from 120 consecutive 


272 


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Not made (moribund) ! Dysentery 








Table II.— continued. 


*78 



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R. S. ...j F. 26 | No No ... ! Mitral rcgurgation Confirmed 






1908 


March April 


Jdy Aag. Sept Oct. 


Nov. 


Dec. 


1909 


50 

45 


Fth. 


May 


Jane 



0 


Chart II.—Notifications of Enteric Fever cases in Kingston during 1908. 



Chart III.—Notifications of En 1 
























































28 


1912 


1913 



Chart VI.— Notifications of Enteric Fever cases during 1912. 



Chart VII.—Notifications of Ent< 






















282 














Chart IX.—Total numbers of notifications of Enteric Fever cases in Kingston, 
month by month, for the years 1908-1914. 













:8 4 



Chart X.—Average of notifications of Enteric Fever cases in Kingston, 
month by month, for the years 1908-1914. 





28s 

ON THE OCCURRENCE AND PRE¬ 
VALENCE OF DISEASES IN BRITISH 
NEW GUINEA 


BY 

A. BREINL 

FROM THE AUSTRALIAN* INSTITUTE OF TROPICAL MEDICINE 


(Received for publication 4 February , 1915) 


Plates XIX—XXVI and Map 

CONTENTS 

Introduction 

PAGE 

285 

Malaria 


288 

Filariasis 


29O 

Leprosy 


*93 

JUXTA-ARTICULAR NoDULES 


*94 

Curious Fevers 


*97 

Yaws 


298 

Venereal Diseases 


*99 

A Peculiar Disease, characterised 

by Arthritis, Osteitis and Periostitis 

300 

Ulcers in the Tropics 


304 

Ulcus Tropicum 


3°6 

Contracting Sore 


309 

Ulcus interdigitale destruens 


313 

Gangosa 

... 

3*4 

Skin Diseases 


3*4 

Concluding Remarks 


3*5 

Appendix I. — List of Papuan Mosquitos 

3*7 

References ... 


3 ** 


INTRODUCTION 

In the following pages are embodied the scientific results of two 
journeys to the coastal belt of British New Guinea, which were 
undertaken for the purpose of mapping out the incidence and 
geographical distribution of tropical diseases among the natives. 

The first journey took place during the months of July and 
August, 1912, and included the coastal belt east of Port Moresby 
and north of Samarai as far as the Mambare River. 


















286 


During the second journey, lasting from the end of June to the 
beginning of October, 1913, the coastal regions west of Port Moresby 
as far as Daru were visited. To the Purari River the journey was 
accomplished on foot, and continued from there in native canoes. 

The coast of New Guinea cast of Port Moresby is fairly densely 
populated, the natives living in numerous villages, ranging in size 
from three or four houses to large settlements with approximately 
3,000 inhabitants, although very large villages are rare. 

The majority of the natives on the south-east coast live by 
agriculture and fishing. The villages consist of a number of houses 
sufficiently large to provide a comfortable shelter for one family. 
Some of the villages are built on high piles in the sea, the houses 
being connected by bridges, but most of the traffic between the houses 
is carried on by native canoes. 

In a great number of the villages situated further inland the 
houses are built on both sides of a main street, since the Government 
advises, for obvious reasons, this scheme of laying out the villages. 
The majority are built on sandy soil and surrounded by more or less 
dense scrub or jungle. 

The coastal districts west of Port Moresby, as far as Orokolo 
Bay, are of the same nature as those on the south-east coast. The 
villages are densely dotted all along the coast, and in many instances 
a few miles only intervene between neighbouring villages. 

After passing Orokolo Bay the nature of the country undergoes 
a complete change. The coast between the Purari and the Fly 
Rivers is broken up into a continuous delta. There are a number of 
large rivers connected by numerous cross channels and cross streams, 
and the land is entirely of a muddy and swampy nature. The 
Villages are built on mud flats, and the traveller may journey for 
days and weeks without stepping on firm soil. 

The villages there consist of houses of varying size; besides small 
huts accommodating one family only, there are very large houses, 
inhabited by a number of families, and large club houses where the 
unmarried boys dwell. 

Throughout the west the native houses are built on high piles, 
and the different sections of each village are connected by bridges, 
consisting of a light scaffolding with loose saplings laid across, which 
are renewed at rare intervals. Still further west, on the Fly River, 



287 


each village consists of a small number of very large houses which 
give shelter to a hundred or more people, and are partitioned off 
into cubicles each occupied by a single family. 

Generally speaking, the natives of the south-east and west coasts 
as far as Orokolo are highly civilized, and have been in constant 
contact and communication with Europeans for nearly a quarter ot 
a century. West of Orokolo the natives become Ids and less 
civilized. In some districts, especially in the neighbourhood of the 
Kikori and Turama Rivers, numbers of villages are not under 
Government control, and many of them have not even been visited 
by Europeans. The villages situated in the Fly Estuary, and the 
inhabitants of some of the islands, as Kiwai Island, are as highly 
civilised as the natives in the neighbourhood of the capital of British 
New Guinea, Port Moresby. 

The medical survey was carried out with all possible thoroughness 
and care, and it may be assumed that cases of almost all diseases 
prevalent in New Guinea have been encountered in one village or 
another. 

During the journey a number of blood specimens were obtained 
in order to map out the distribution of filariasis and malaria. 

The less civilised the districts, the fewer natives were found 
willing to supply a drop of blood for a film. Unfortunately in 
several districts, especially around the Kikori River, it was 
impossible even to obtain photographs of natives suffering from the 
various complaints. As opportunity offered, sores were scraped with 
a sharp spoon, and the granulation tissue so obtained preserved in 
70 per cent, alcohol for future microscopical examination. 

Speaking generally, a number of diseases such as various types 
of ulcers, malaria, filariasis and elephantiasis, juxta-articular 
nodules, gangosa, yaws, and several skin diseases, mainly tinea 
imbricata, were encountered in most of the villages visited. It must 
be borne in mind that for a considerable time frequent intercourse 
and trade took place amongst the natives on the south coast between 
Orokolo and Samarai. Every year the natives of Port Moresby and 
several adjoining villages, who are the potters of New Guinea, travel 
west in the lakatois (large sailing canoes), exchanging their pots for 
sago which grows abundantly in the western parts. The journeys 
must have offered ample opportunity for the spread of contagious 



288 


and infectious diseases. In more recent times the opening up of the 
couhtry, and the drawing of the supply of native labour from 
districts the inhabitants of which had hitherto not been wont to 
travel, has given rise to new channels for the propagation of disease. 
It is, however, noteworthy that several diseases, such as gangosa, 
juxta-articular nodules, yaws and tinea imbricata, and different 
types of sores, were seen in natives who were unable to travel any 
considerable distance without endangering life and limb on account 
of hostile neighbours. 


MALARIA 

With the exception of a few districts, malaria occurs with 
varying severity throughout British New Guinea, although its 
frequency is dependent on seasons and local conditions. New 
Guinea, situated within the monsoonal belt, has two distinct seasons 
—the north-west beginning at the end of November and lasting 
until March or April, when land breezes prevail, and the south-east 
season when the prevailing wind comes from the sea. 

In the coastal villages malarial fever is more prevalent during 
the north-west season when mosquitoes are in evidence. During the 
south-west time, on the other hand, malaria is far less common, since 
the sea breeze prevents the mosquitoes from swarming into the 
villages. 

In the inland villages in sheltered positions, on the whole, no 
marked seasonal variation is observed. Coastal villages in exposed 
positions are comparatively free from fever, whereas coastal villages 
sheltered from the breeze show a large percentage of children with 
enlarged and palpable spleens. In the villages west of Orokolo, 
built on mud flats, which were visited during the south-west monsoon, 
no cases of malarial fever amongst the natives were encountered, 
although fever is said to be prevalent during the north-west season. 

The natives contract malaria as a rule in early childhood, and 
acquire, in the course of years, an active immunity against the type 
of malarial infection prevalent in their place of residence. The 
incidence of malaria in natives decreases in direct proportion to their 
age. 

The number of children with enlarged spleen gives a definite 



289 


indication of the amount of malaria in a given locality. The spleen 
census was taken by determining the number of children with 
enlarged spleens in the different villages, and blood films were made 
of those children who showed considerable enlargement of the spleen 
to ascertain the type of malaria prevalent. 

On several occasions the spleen index of babies in arms was taken 
separately from that of children of ages ranging between 5 and 11 
years, and the former was considerably higher than the latter, in 
one instance 95*1 in comparison with 58*6. 

The accompanying map shows that simple and malignant tertian 
types of malaria are fairly equally distributed and more common 
than the quartan type; the latter is more prevalent in the furthest 
north-east corner, on the mouth of the Mambare River, and is not 
infrequently met with in the Mekeo district. 

Out of 245 blood films taken from a small number of native 
children, 132 contained malarial parasites, sometimes in considerable 
numbers; at other times only a few ring forms were seen in a whole 
film. 

Out of 245 blood films examined, 48 contained the parasite of 
malignant tertian, 34 of simple tertian, and 27 of quartan malaria. 
In 23 the type of the parasite could not be determined with certainty. 

The white population of British New Guinea suffers, with hardly 
any exception, from occasional attacks of malarial fever, and not one 
white settler was met who had been entirely free from attacks of fever 
during his stay in the country. 

Of Anophelinae, N yssorhynchus annulipes and Cellia punctulata 
were collected, the former being the more common mosquito, 
occurring in the proportion of about 10 to 1 of the latter. 

The occurrence of A 1 yssorhynchus annulipes in numbers through¬ 
out the districts where malarial fever is most prevalent, and the fact 
that the development of Plasmodium falciparum has been traced by 
Kinoshita (1906) in this mosquito, make it extremely likely that 
Nyssorhynchus annulipes acts as intermediary host of the malarial 
parasite in New Guinea. 

Of other Anophelinae, N yssorhynchus bancrofti has been 
collected, but this species does not transmit the parasite. 



290 


FILARIASIS 

Thorpe in 1896 drew attention to the fact that the microfilaria 
in the blood of the natives of the Tonga Islands does not show the 
same periodicity as observed elsewhere. Lynch (1905) confirmed 
this statement, finding that in 105 out of 156 cases of filariasis 
amongst} the Fijians, microfilariae were present in the peripheral 
blood both by day and night. 

Similar observations in the cases of filariasis among the natives 
of the Philippine Islands led Ashbum and Craig (1907) to the 
creation of a new species, Filaria philippensis. The larvae, although 
resembling closely Microfilaria nocturna , do not possess any 
periodicity, and occur in approximately the same numbers in the 
peripheral blood during all hours of the night and day. The 
movements of this microfilaria are described as progressive and 
lashing, the sheath is very tightly fitting, so that the microfilaria 
cannot slip backwards and forwards within it, as in the case of 
Microfilaria nocturna. At the posterior end the tail diminishes 
progressively and uniformly to a fine thread-like point. Bahr (1912), 
Fiilleborn (1913), and many other observers, have failed to 
discriminate between the larvae of Filaria philippensis and that of 
Filaria bancrofti. 

Fiilleborn (1911), utilising material collected in the Bismark 
Archipelago and German New Guinea, pointed out that 10 to 60 
per cent, of the natives showed large sheathed microfilariae in their 
blood, which were of the same anatomical structure as Microfilaria 
bancrofti but did not possess the typical nocturnal periodicity. More¬ 
over, the adult worm of the Fijian microfilaria, according to Leiper, 
cannot be differentiated from Filaria bancrofti. 

Fiilleborn (1912) concluded, therefore, that in the present state 
of our knowledge two biological varieties of Filaria bancrofti can be 
differentiated; the larvae of one variety have a pronounced nocturnal 
periodicity, whereas the larvae of the other, mostly occurring in the 
Pacific, are present in the peripheral blood at all hours of the day 
and night. 

A troop of twenty-three Samoans, who were visiting Europe, were 
examined by Fiilleborn (1912). He found that nearly half of them 
harboured microfilariae in their peripheral blood, morphologically 



291 

identical with Microfilaria nocturna , but without any marked 
periodicity. 

Whilst travelling through the coastal districts of British New 
Guinea, thin smears were made from the day blood of a number of 
men, women and children to be examined for the presence of micro¬ 
filariae. It was found impossible to use thick films for this work, 
since previous experience had shown that they deteriorate much 
quicker than thin blood smears, and after exposure for some time to 
the moist tropical heat are difficult to dehaemoglobinize. On account 
of the short stay in the different villages it was possible to take 
blood films in the day-time only, as the blood examination of the 
same natives on two occasions would have meant too great a delay. 
A more thorough and extensive filarial survey will be undertaken at 
a future date. 

Although the number of blood examinations was comparatively 
small, the results indicate that the natives living in the coastal belt 
of eastern and north-eastern New Guinea harbour filarial larvae in 
their blood to a greater extent than in the western parts. Out of 166 
blood slides taken at random on the first journey, twenty-four 
contained microfilariae, giving an infection index of slightly over 
17 per cent., whereas out of 166 films taken during the second 
journey, microfilariae were discovered in eight slides, being about 
5 per cent. A more extensive survey, taking samples both in the day 
and night time, would almost certainly reveal a much larger 
percentage of infection. 

On two occasions only the blood of some of the carriers was 
examined, and two and three respectively, out of about fifteen, 
proved to be infected with filarial larvae, which showed the typical 
nocturnal periodicity. 

Morphologically the microfilaria found in the blood of New 
Guinea natives does not differ essentially from typical Microfilaria 
nocturna. Table 1 gives the measurements of a number of micro¬ 
filariae, stained by Giemsa's method, and for comparison are 
appended Fiilleborn’s average measurements of typical Microfilaria 
nocturna and of microfilariae found in the blood of Samoans. 

The experience gained in New Guinea coincides with Bahr's 
observations in Fiji, and Fiilleborn's in the South Pacific, that there 
exists in British New Guinea a microfilaria morphologically identical 



292 


Table I.—Measurements of Microfilariae found in the blood of New Guinea natives; films taken 
in the daytime. 



Nerve spot Excretory 1 
from j pore from : 

anterior end anterior end 

Inner body 
from 

anterior end 

1 

Length of 
inner body 

j Gi cell 
' from 

' anterior end 

| Anal pore 
from 

! anterior end 

1 

Total 
j length 


% 

0 

,0 

% 


0/ 

,0 

I 0/ 

/o 


1 

! * 8-5 

28*1 

468 

5 ° 

... 

81*4 

4<>5 

2 

1 9 * 1 

28-5 

52-3 

5 ° 

73*8 

88 -o 

410 

3 1 

* 7*3 

27-9 

486 

38 


848 

358 

4 ! 

1 6* > 

26-6 

48-9 

60 

689 

85*3 

450 

5 ; 

20-1 

* ... 

57*4 

35 

74 * 1 

... 

340 

6 1 

> 9*3 

286 

45*7 

7 ° 

73*4 


35 ° 

7 ! 

1 

19*6 

29*4 j 

40*0 

60 


86-8 

3<>5 

8 

1 

1 6*4 

26*2 

47*8 

55 

73*8 

« 7'3 

366 

9 

18*2 

27*3 

50-0 

45 

7**5 ; 

82*9 

440 

1 ° j 

21 *o 

29-5 

5 1 * 1 

52 

74*5 

88-0 

1 352 

.. ! 

1 

19-6 

316 

607 

30 

74*4 

88*5 

395 

12 ! 

■S -5 

3°*3 

48*4 

60 

75*7 


33 ° 

*3 


28-0 

49*6 

56 


87*0 

3 *° 

<4 

i »-4 

29-0 

5°*4 

54 

7 ! ‘4 


245 

Average 18-7 

! *8-5 

49*9 

5 * 

73 *» 

860 

362 

Fulleborn’s average measurements of typical Microfilaria nocturna — 




iR -3 

j 28*6 



70*6 

1 

83-5 : 

232*5 

Average of non 

•periodical Samoan Microfilaria- 

- 





19*3 

1 1 

1 285 

... 


| 1 

70*2 

82*9 | 

272*5 



*93 


with Microfilaria nocturna , but not possessing the typical periodicity, 
in addition to microfilariae which are only present in the peripheral 
blood at night time. 

It is interesting to note that only one specimen of Stegomyia 
pseudoscutellaris was collected in Samarai on the south-east coast 
of New Guinea, which species was not encountered anywhere else. 

The patchy distribution of filariasis is quite as well marked in 
New Guinea as in other parts of the tropics; for example, in one 
village nearly 3 to 5 per cent, of the male population suffered from a 
soft swelling of the groin glands, so typical in filariasis, whereas in 
the neighbouring villages cases were only rarely noticed. 

Cases of elephantiasis were seen in varying numbers throughout 
the districts visited. Of nearly every case of elephantiasis blood 
films were taken, but in no instance were microfilariae found in the 
blood. The arms and legs were most frequently affected, the arms 
perhaps more often than the legs. The genitalia were not so 
frequently implicated. 

Plate XIX, fig. 1, shows a typical and far advanced case of 
elephantiasis of the scrotum, affecting at the same time one leg; and 
fig. 2 shows a typical case of elephantiasis of the labium minor of the 
vulva. 


LEPROSY 

The distribution of leprosy in New Guinea is of some interest. 
This disease has already been referred to by W. MacGregor in the 
yearly report for New Guinea, 1888-1889, where it is stated that 
‘ Leprosy is not an uncommon disease amongst the natives, but does 
not present its worst form and has not been communicated to any 
white person.' 

On the whole, in the districts visited, leprosy was found endemic 
in only three, namely, the Trobriand Islands—a number of islands 
off the north-east coast—the Mekeo district on the St. Joseph River 
in the Central Division of British New Guinea, and in one district 
of the Gulf division. 

Only very few cases were seen on the south coast east of Port 
Moresby, and not one definite case west of the Vailala River. 
Altogether eighteen cases were diagnosed, four cases in the 
Trobriand Islands, seven in the Mekeo district, and eleven in the 



villages on the south coast between Isiu and Orokolo, west of Port 
Moresby. 

The three different types, namely, nodular, nerve, and mixed 
leprosy, were found. 

Plate XIX, fig. 3, shows one of the most advanced cases of 
leprosy seen. 

It is an interesting fact that the natives of the Trobriand Islands 
know leprosy well, having a special native name for the disease, 
namely, Kai-gwai-guia. When the natives were asked to bring 
forward these cases, all produced were typical lepers. 

Considering the number of natives seen, and the large area 
traversed, if becomes apparent that leprosy is present in New Guinea, 
although by no means prevalent. A great number of ulcers of various 
descriptions resembled leprosy at the first glance, but careful 
examination excluded the diagnosis. 

JUXTA-ARTICULAR NODULES 

The occurrence of juxta-articular nodules was first described by 
MacGregor in 1901 in New Guinea. Later, in 1904, Steiner described 
the occurrence of tumours amongst Javanese, appearing as hard, 
round irregular nodules below the skin, and situated always in the 
neighbourhood of the joints. The skin above the tumours was 
moveable, and the nodules were not connected with the underlying 
fascia. On histological examination these nodules consisted of 
concentric layers of hard fibrous tissue, which was degenerated in 
the centre into coarse, irregular, structureless masses. Various 
staining methods did not reveal the presence of any bacteria. 

Two years later, Jeanselme (1906) described the occurrence of 
similar tumours amongst the natives of Siam. Microscopic examina¬ 
tion showed that they were formed of three layers—a central one 
consisting of degenerated tissue, an outer zone of inflammatory 
reaction, and an intermediate or transitional zone. 

Gros (1907) observed the occurrence of similar tumours amongst 
the Algerian natives in small numbers only; ten cases were seen out 
of approximately 12,000 patients. Histologically they were of 
similar structure to those examined by Jeanselme. 

The occurrence of the same disease was reported later by Neveux 



2 95 


(1907) amongst the natives of Senegambia, and Fontoynont and 
Carougeau (1908) made an extensive study of this complaint in 
Madagascar. In their experience, juxta-articular nodules were 
found in persons of both sexes and of all ages, though the male sex 
was more frequently attacked by the disease. Special attention was 
drawn to the symmetrical occurrence of these nodules. They 
pointed out that the seat of predilection of the tumours was the outer 
surface of the extremities, especially the extensor surfaces, also, that 
they occurred in groups in the skin in the neighbourhood of joints, 
and especially where bone lies immediately below the skin, as near 
the elbow and large trochanter. 

The development of the juxta-articular nodules is very slow, 
extending over years, until finally they become fibrous or fibro¬ 
cartilaginous. Fontoynont and Carougeau found in early nodules, 
amongst the soft caseous central mass, very small white granules, 
consisting of mycelium and filaments of a fungus, which they 
described and termed Discomyces carougeaui . 

Inoculation of this fungus into monkeys, rabbits, and guinea- 
pigs, gave no results. 

Lebouef (1911) observed four cases of this disease in New 
Caledonia. According to Joyeux (1913) similar tumours were 
frequently seen in natives of French Guiana, though the microscopical 
picture differed in some essential points from that given by 
Fontoynont and Carougeau. These tumours consisted of a ground 
substance of fibrous tissue contained within inflammatory foci, 
mostly around the blood-vessels. The absence of bacteria in the 
sections caused Joyeux to consider juxta-articular nodules as a 
clinical syndrome resulting from widely different causes. 

Juxta-articular nodules are of frequent occurrence throughout the 
coastal districts of British New Guinea. In a great number of 
villages many cases were seen, only a few in others. The clinical 
manifestation did not differ from that described by previous 
observers. Fibrous tumours of varying size, from a pigeon’s to a 
hen’s egg, occurred in different parts of the body, mainly in the 
neighbourhood of joints and most frequently near the ankle joints 
(fig. 4) above the large trochanter and over the olecranon (Plate XX, 

ftg- 5 )- 

The subcutaneous tumours were round or oval, the skin above 



296 

freely moveable, and they were not connected with the underlying 
muscle. 

They were of hard consistence, having as a rule a smooth, rarely 
an irregular, surface, and occurred in the majority of cases 
bi-symmetrically. In no cases were fistulous openings observed. 

The majority of the affected natives were middle-aged men, only 
rarely women. One young girl, about 16 years of age, showed well- 
developed nodules over both elbow joints. 

In some villages situated in the eastern end of British New 
Guinea as many as 7 to 10 per cent, of the whole population were 
affected. In other villages closely adjoining even the most pains¬ 
taking search did not reveal the existence of a single case. 

These tumours never caused the natives any pain or inconvenience, 
and it was only after a great deal of persuasion that permission was 
given to extirpate one of these tumours for histological examination. 
Further material was received through the kindness of Dr. Giblin 
in Samarai.* 

Macroscopically, in cross sections, the nodule does not differ in 
any respect from a subcutaneous fibroma; it consists of strands of 
connective tissue, concentrically arranged, the skin above being not 
connected with the tumour, which is apparently not encapsulated. 
Here and there are small irregular areas of yellowish appearance, 
showing softening. 

On microscopical examination of three nodules, the three zones 
described by the previous observers could not be distinguished. 

One of the nodules, which was extirpated with the overlapping 
skin, showed that the nodule itself was well defined from the 
surrounding subcutaneous tissue, which was normal except for the 
presence of a certain amount of small-celled infiltration and 
hyperaemia. 

The nodule itself consisted of dense concentric layers of 
connective tissue cells, possessing large irregular nuclei and cyto¬ 
plasm with fibrillar-like processes, which anastomosed and formed 
an open meshwork. Here and there, irregularly distributed 
throughout the nodule, were small areas where the cell nuclei had 
disappeared, the tissue still showing the original texture. Numbers 

• Dr. Giblin ha» informed me since that he had treated one or two Europeans suffering from 
juxta-articular nodules. 


297 


of leucocytes were scattered throughout, but were most numerous in 
the degenerated parts. On the whole, the tissue of the nodules was 
well vascularised, and there was small-celled infiltration around the 
blood vessels. 

The bacteriological examination of the sections did not show any 
microorganisms, and none of the nodules showed the ‘ Dtscomyces 
carougeaui * It is probable that the fungus may only be present in 
the early stages, and that the specimens obtained were of cases of 
long duration. 

CURIOUS FEVERS 

The short stay in the different villages did not afford much 
opportunity for studying the different types of fevers prevalent. 

Four cases of a curious and apparently specific fever were 
observed in villages on the south-east coast. All four cases showed 
the same clinical symptoms, which were sufficiently pronounced to 
suggest a specific fever. 

According to an account given by an employer of native labour, 
the disease sets in gradually, with an irregular fever. The first 
symptom noticeable is an icteric discoloration of the mucous 
membranes and of the sclerae, accompanied by pain in the splenic 
region. The patients suffer from more or less frequent attacks of 
continuous high fever, which may last for a few days or even a week, 
the temperature rising to 105° F. They emaciate considerably. 
The disease runs a chronic course—one of the patients had been ill 
for nearly two and a half years—improvement and relapses following 
each other at irregular intervals. The patients die finally with 
symptoms of great emaciation and pronounced icterus. 

The examination of the four cases showed a slight enlargement 
of the spleen and a very marked enlargement of the liver, which in 
one case extended to five finger breadths below the ribs; the sclerae 
and the mucous membranes were of a dark yellowish colour. The 
blood was decidedly anaemic. 

The disease is well known to the natives, and is called ‘ Gobora * 
in the Mailu district, and ‘ Tebi* further east. 

Blood films were taken of each of the cases. Two of them showed 
a slight malaria infection, whilst in the others no malaria parasites 
could be detected. No other parasites were seen in any of the blood 
films. 



298 


YAWS 

Throughout the coastal districts of British New Guinea yaws is 
widely distributed, and barely a single village was found entirely 
free from the disease. In some districts yaws is more prevalent than 
in others. 

In 1897 Sir William MacGregor drew attention to the fact that 
‘Yaws is universally endemic, but it is greatly milder than it is in 
the Pacific, and is in fact not much taken note of. There does not 
even appear to be a native name for it in any dialect known. No 
European appears to have contracted this disease in the Possession. 1 

The majority of the sufferers were children, and all the typical 
lesions of primary and secondary yaws were seen at one time or 
another. It was noteworthy that in different districts one or the 
other of the clinical types of yaws prevailed. In the village of 
Orokolo, for example, many of the children examined suffered from 
the typical granulomatous eruption around the anus, whereas in 
other districts hardly one case with this manifestation was seen 
amongst fifty children suffering from yaws. 

In the western parts sporadic cases of a generalized eruption of 
secondary yaws were observed. The sufferers were mostly children, 
and clearly showed that secondary yaws might lead to considerable 
destruction of the skin. In one case, a young girl (figs. 6 and 7), 
the eruption had implicated the nose, both cheeks, lip, the upper 
arm and elbow, and had formed a semicircle on chest and back. 
The arm was contracted at the elbow joint. On the affected parts 
prominent yellowish scales were seen, which covered ulcers, roundish 
in the early stages and later confluent. The fundus was formed by 
raised reddish granulation tissue. In between these scabs was, here 
and there, newly formed glossy skin. 

In another case (fig. 8), the nose was partly destroyed by the 
specific granuloma, which spread from the nose to the cheek. 

Yaws was as prevalent amongst the civilised natives as amongst 
those who had hardly come under the Government influence, and 
had had no intercommunication with the neighbouring tribes. 



VENEREAL DISEASES 


It is very difficult indeed to map out the distribution of venereal 
disease amongst natives during a journey from village to village. 
Natives are prone to hide venereal disease from each other, and 
systematic inspection, especially of women, is practically impossible. 
The diagnosis of ‘ Syphilis 9 in a native community is more difficult 
than in Europeans, on account of the occurrence of various ulcers 
and after effects of ulcers, which closely resemble the syphilitic 
lesions. Many of the manifestations of the tertiary or late stage of 
yaws, as described by Castellani, are identical with those of tertiary 
syphilis. 

From the limited observations made, syphilis has not spread to 
any extent amongst the natives in the coastal districts of British 
New Guinea. This is in all probability due to the strict measures 
adopted by the Government against its spread. Lock hospitals 
have been established in different districts, and all cases reported are 
interned and only discharged after thorough treatment. 

A scanty number of isolated cases were found here and there, 
but only in those districts where the natives had come in close contact 
with Europeans. 

ULCERATIVE GRANULOMA 

A few cases of ulcerative granuloma were seen in the eastern 
parts of New Guinea, several of them under the care of Dr. Giblin 
in the Lock Hospital at Samarai. Only three typical cases were 
encountered in natives living in villages on the east coast, and a few 
sporadic cases in the western districts. 

GONORRHOEA 

Cases of gonorrhoea may be found in all localities where the 
natives have come in contact with Europeans. In several villages in 
the Goaribari district, where communism is still existent, from a 
third to a half of the population were suffering from the typical 
lesions due to Gonococcus. The clinical symptoms did not differ in 
severity from those observed in a temperate climate, although 
venereal sores were more frequent on account of the lack of personal 
cleanliness. 



300 


A PECULIAR DISEASE CHARACTERISED BY ARTHRITIS, 
OSTEITIS AND PERIOSTITIS AMONGST NEW GUINEA 

NATIVES 

In his article on skin disease in the tropics, Plehn (1914) mentions 
that Castellani, Howard, Schiiffner and others have attributed joint 
affections and periostitis to yaws, regarding them as its tertiary 
stage. He also states that he himself observed similar bone lesions 
in Cameroon natives, which he considered as the final result of 
chronic osteomyelitis. Skiagrams, appended to the publication, 
show thickening of the periosteum of the bones of arm and hand in 
children. 

A disease characterised by similar symptoms was encountered 
amongst New Guinea natives of different ages, of whom, in several 
instances, fairly reliable histories could be obtained. 

The distribution of these cases was irregular, many villages being 
quite free, whilst others furnished six or more cases, most of which 
failed to show any signs of previous attacks of yaws. No evidence 
could be gathered as to a connection between the incidence of these 
two diseases. 

It would be premature to state or deny definitely any relationship 
between yaws and the disease in question, until cases have been 
observed, where the two merge into each other without any prolonged 
interval. The irregular distribution, the similarity of the clinical 
histories, its absence from tropical regions where framboesia is very 
prevalent, are points in favour of the supposition that the two 
complaints are separate clinical entities. 

This disease is exceedingly chronic, and is termed by the natives 
of the central district ‘ Roaki,* and by the natives of Kiwai Island 
‘ Buuo* or ‘Auma.’ 

The most prominent symptom is an affection of one or more of 
the joints, ranging from a hot painful swelling to suppuration and 
the formation of sores with fistulae in the skin above the joints, 
which discharge an amber-coloured clear liquid containing floccular 
particles. At the same time spindle-shaped thickenings of the larger 
and smaller bones of the extremities become noticeable, which, 
especially when the small bones of the hand and foot are implicated, 
may lead to softening and resorption of the bone. Similar sores to 



3oi 


those above the joints may make their appearance above the 
swellings of the bone. 

The disease affects natives of all ages, men, women and children, 
although the majority were young persons. The youngest patient 
was about eight years of age. 

As far as could be ascertained, the disease begins with an 
affection of one or more joints, without any special predilection being 
noticed. The joints swell up and become painful. After a varying 
period the swelling may abate or increase, and in this case the pain 
prevents the patient from using the affected extremity. After a 
prolonged period fistulous openings may appear in the neighbour¬ 
hood of the joints secreting at first flocculent pus, and later on an 
amber-coloured clear fluid. 

The foot so affected (Plate XXI, fig. 9) closely resembles Madura 
foot without the presence of the typical granules in the pus. 

In other cases the bones may become affected, at first either in 
close proximity to the joint or the diaphysis of the long bones, mostly 
the tibia, ulna or radius; or in still other cases the metatarsal and 
metaphalangeal bones may show the change. The bone thickens, 
and a spindle-shaped swelling becomes noticeable which is not 
painful on pressure. This thickening is so well marked that it can 
readily be noticed on inspection (see figs. 10, 11). 

These swellings of the various bones may remain unaltered for 
years, even for the rest of the patient’s life-time. In other cases, 
especially when the smaller bones of hand and foot are the seat of 
the swelling, the lesion clinically resembles osteomyelitis. Fistulous 
openings appear which discharge copiously at first, and in time the 
whole of the affected bone may become resorbed. This process leads 
to peculiar disfigurement of the feet and hands (fig. 12). In cases 
where one of the metacarpal or metatarsal bones is resorbed, the 
fingers or toes become either retracted or even displaced, as in 
Plate XXII, fig. 13, where the large toe is in an abducted and hyper- 
extended position. 

The frequent occurrence of the swelling of the radial, ulnar and 
tibial bones simultaneously with swellings of the metacarpi or meta¬ 
tarsi and swellings of the joints, appeared to furnish proof that all 
these lesions were due to the same disease. 

The lesions of the hand and feet referred to may easily give rise 



302 


to a mistaken diagnosis. More than one case on superficial 
examination resembled nerve leprosy. A thorough examination, 
however, excluded this diagnosis, since in every case the finger tips 
were normal, the nails still well formed, and the thickening of the 
main nerve trunks was absent. 

In a later stage ulcers are formed in the skin over the affected 
joints and bones. These are superficial, of varying size up to 
25 cm. in length, oblong in shape with irregular edges; the granula¬ 
tion tissue is only slightly raised, of dark reddish colour, discharging 
an amber-coloured clear fluid. On the whole they show a tendency 
to spontaneous healing, forming dense scar tissue (fig. 14). 

It was impossible to obtain any conclusive evidence as to the 
sequence of the lesions of bone and skin, since the information 
obtained from intelligent natives varied considerably. Some of them 
pointed out that the bone and skin lesions arose independently, 
others, however, stated definitely that the bone lesion preceded the 
skin lesion. 

Several of the cases resembled, clinically, osteomyelitis with 
fistulous openings. In the majority, however, an inserted probe 
did not reach the bone. 

The histological examination of the granulation tissue of one case 
did not show any peculiarities. It resembled, microscopically, typical 
granulation tissue containing a number of phagocytes and numerous 
bacteria and cocci on the surface. 

Short case histories so far as obtained, and a clinical description 
of a few typical cases, may be helpful to illustrate this interesting 
and somewhat obscure disease. 

Case /. Idiri—Woman about 35 years of age. 

The patient had been ill for a long time, perhaps five years. She 
never suffered from any painful joint lesions. The first symptom 
noticed was a hard swelling of the tibia. A short time afterwards 
sores formed above the swelling of the bones, which healed up after 
a long time. The metatarsus of the big toe of her right foot began 
to swell, and afterwards small openings appeared in the skin above 
the swelling, discharging, for a long time, thick matter. A 
considerable time afterwards these fistulae ceased to discharge, and 
healed up with displacement of the large toe. 



303 


At the time of examination the shaft of the right tibia showed a 
considerable spindle-shaped swelling. Numerous scars had formed 
in the skin above and on the dorsal side of the foot. The metatarsus 
of the big toe had disappeared, and the toe consequently was 
abducted and hyperextended. The right wrist joint was swollen 
and painful, the ulna showed in the middle of its shaft a marked 
spindle-shaped swelling, about io cm. long. 

Case IL Boy about 12 years of age. Comp. fig. 11. 

The boy had been ill since childhood, and the disease had begun 
with painful swellings of the bone. Some time afterwards large, 
slowly healing sores formed in the skin above the swellings. 

The boy was very emaciated. Both tibiae were curved forward, 
the bone thickened and the skin above showed ulceration. Both 
ulnar bones had, in their upper third, a spindle-shaped thickening 
with a fistulous opening, on one side discharging liquid pus. There 
was extensive ulceration on both the arms, where roundish irregular 
islets of reddish, raised granulation tissue were surrounded by very 
thin newly-formed skin. Both legs were ankylosed at the knee 
joint. 

Case III . Small boy about 8 years of age. 

The patient was hale and hearty till about one year ago, when 
his right ankle joint began to swell up. Shortly afterwards his right 
knee joint became painful and swollen, and later the left knee and 
both wrist joints became affected in the same way. When examined 
the boy was considerably emaciated, both the knee and wrist joints 
were swollen and painful. The muscles of both calves as well as the 
muscles of both forearms were atrophic. The distal end of both of 
the radial bones was considerably enlarged. 

The patient did not show any sign of present or past infection 
with yaws, and throughout the village not a single case of yaws 
was seen. 

Case IV. Man about 35 years of age. 

According to information obtained from the patient, the disease 
had started about twenty years ago with pains in his back and in 
the joints of his arms and legs. 

When examined his right foot was very much swollen, and all 



304 


the toes of his foot had dropped off as the result of ulceration. The 
skin of the foot was uneven, lumpy, and numerous fistulous openings 
were seen, some apparently closed, others secreting on pressure 
whitish floccular masses. The muscles of the leg itself were atrophic, 
and the skin above the much thickened tibiae covered with old scars. 
The left tibia showed the same changes, simulating a chronic 
oestomyelitis. The left elbow joint was swollen, hot and painful, 
and the tip of the elbow the seat of a small ulceration. 

The fingers of the left hand were deformed, the metacarpo¬ 
phalangeal joint of the fourth and fifth finger were ankylosed. The 
shafts of both radial bones showed a spindle-shaped thickening, and 
there were deep scars over both radio-carpal joints. 

ULCERS IN THE TROPICS 

A great variety of superficial and phagedaenic ulcers, apparently 
of the most varied etiologies, are collected under the name 4 Tropical 
Ulcer.’ The clinical differentiation of ulcers in the tropics becomes 
still more involved on taking into consideration the careless habits 
and the lack of cleanliness amongst native races in general. Any 
superficial skin abrasion on hand or leg, which if at all cared for 
would heal, may give rise to a small ulcer which on account of the 
continuous irritation caused by sand, dirt and flies may spread. It 
is probable that many of the large phagedaenic ulcers are not due to 
one specific microorganism, but are caused by a variety of micro¬ 
organisms invading a small wound due to an accident. 

The observations made in British New Guinea appear to bear 
out this statement. On the whole the incidence of ulcers depends 
largely on the state of cleanliness, which is greatly influenced by the 
natural conditions and surroundings of the different villages. In 
the districts west of Orokolo, where the villages are built on high 
piles on muddy grounds, and where fresh water is very scarce, 
ulcers of varying types are encountered in great numbers among the 
natives. 

The natives in these districts, where sago palms grow in great 
abundance, use sago as their staple article of diet, and spend, in 
consequence, much of their time in preparing sago from the trunk 
of the palm. The sago palm carries large thorns on the rib of the 



305 

leaves and on the trunk, and the natives are often injured by them 
when felling the trees. The wounds in the majority of cases are not 
cared for, and undoubtedly many ulcers are originally due to wounds 
caused by these thorns. 

The number of inhabitants with ulcers in villages built on dry, 
sandy soil is much smaller. 

The hypothesis that many of these ulcers are caused through 
secondary contamination of smaller or larger injuries of the skin 
receives additional support from the experience gained in their treat¬ 
ment. A great number of the sores heal up readily under the 
application of antiseptic powders only, without any specific 
treatment. It must, however, be, kept in mind that the scar, 
covered by a newly-formed skin on the original site of the ulcer, is 
always more liable to injury and may again become the seat of a 
similar ulcer. 

The question of the differentiation and diagnosis of ulcers 
becomes more complicated by the facts that in the majority of the 
natives the ulcers are in an advanced stage, that it is impossible to 
obtain a definite and reliable history, and that advanced ulcers of 
many different etiologies are similar. The question of a tertiary 
form of yaws is still under discussion, and the clinical symptoms 
attributed by various observers to tertiary yaws may be due to other 
factors. 

The diagnosis of syphilis is not less complicated, and a traveller 
with little experience of natives in their original state might make 
the diagnosis of syphilis where its occurrence may be excluded with 
certainty. 

Gangosa, so similar to tertiary syphilis in its advanced stages, 
causes ulcers of arms and legs which, when far advanced, are in no 
way characteristic. 

Many other complaints, such as leprosy, various skin diseases, 
etc., may take an atypical form in natives and make a differential 
diagnosis in the late stages impossible. 

Histological examination of the granulation tissue does not 
permit a definite differentiation of the ulcers, since the microscopic 
appearance of sections of different and clinically apparently well 
characterised sores is similar. Smears taken from the surface are, 
as a rule, a pell-mell of saprophytic bacteria and yeasts of various 



306 


sizes and shapes, whilst smears taken from the deeper parts 
generally do not show any bacteria. 

The examination of the natives of the south coast of British New 
Guinea revealed the presence of a great number of ulcers. Several 
were well characterised and could be classified, whilst others did not 
show any typical features. 

TROPICAI, SLOUGHING PHAGEDAENA, ULCUS TROPICUM 

Ulcus tropicum, a complaint occurring in practically every part 
of the tropics, is known under nearly as many names as there are 
localities where it has been found. 

Clinically it is a chronic sloughing ulceration of phagedaenic 
character, which may spread into the depth, laying bare muscles and 
bones and showing a tendency to spontaneous healing, or leading, 
if neglected, to considerable deformities of the affected extremities. 

The literature on tropical sore is fairly extensive, and has been 
summarised by Plehn (1914). It is, however, beyond doubt that 
several varieties of sores, specific and non-specific, have been 
previously included under the same heading. 

Prowazek (1907) described a spirochaete, Spirochaeta schaudinni , 
as the specific cause of the disease. This spirochaete is actively 
motile, resembling the balanitis spirochaete, possessing only rarely 
periplast on one end; it multiplies by longitudinal fission and forms 
involution forms. 

The same spirochaete was found by Keysselitz and Mayer (1909) 
in smears taken from the deep layers of ulcers, and by Wolbach and 
Todd (1912); although the latter authors consider that 'The fact 
that these ulcers are almost invariably found solitary and on regions 
exposed to trauma supports the explanation that the process is a 
reaction to an organism having slight power of invasion.’ 

Le Dantec (1911) had previously pointed out that two conditions 
were essential for the development of a tropical sore, namely an 
abrasion of the skin and exposure of this to soiling by humid earth. 

A direct experimental transmission of the sore to animals such 
as guinea-pigs, monkeys, and apes and to men has been 
unsuccessful in the hands of all observers. Blanchard (1914) 
succeeded in transmitting a tropical ulcer from man to man, but 



307 


only after having produced at first an artificial necrosis of the skin, 
which was some days afterwards inoculated with the granulation 
tissue from a case of tropical ulcer. 

Cases of tropical ulcers were found in varying numbers, 
throughout the coastal districts of British New Guinea, and barely 
one village was entirely free from this complaint. 

In the villages situated on high, dry ground only a small number 
of cases was seen, whereas in the villages in the western parts of 
New Guinea, which are built on swampy, muddy ground, a great 
number of cases was encountered. It seems, therefore, that the 
frequency of tropical ulcers stands in some relation to the incidence 
of mangrove mud. 

The tropical ulcer of New Guinea does not differ essentially from 
that observed in other parts of the tropics. It occurs mostly on the 
lower extremities, on the dorsal surface of the foot, in the neighbour¬ 
hood of the external and internal malleolus, on the skin, rarely on 
the back of the thigh. In the early stages the ulcer resembles a boil. 
There is an ulcer in the centre which is covered by a scab, slightly 
raised above the surface of the skin, showing fluctuation and a 
certain amount of inflammatory reaction around. Very soon, if 
neglected, this small round ulcer begins to spread, the epithelial 
layer breaks down, showing underneath a fungating ulcer, which 
bleeds very readily and is covered with a small amount of thick 
greyish pus. The ulcer is well defined from the surrounding tissues. 

In the beginning the ulcer spreads very rapidly, is irregular in 
outline, and covered by a layer of foul smelling greyish semi-fluid 
pus, which forms a pseudo-membrane covering the dirty greyish to 
reddish granulations (fig. 15). After a varying interval the ulcer 
spreads into the depths, destroying the subcutaneous tissue and the 
muscles (fig. 16) and often attacking the underlying bone, and gives 
rise to periostitis and osteitis. Now and again a true osteomyelitis 
may follow the ulcer, and then pieces of necrotic bone are often found 
embedded in the granulation tissue. In other cases, however, the 
ulcer remains superficial and the granulation tissue proliferates and 
projects above the level of the surrounding skin. The borders in 
chronic ulcers are either sharply defined or sometimes undermined. 
The whole of the foot may swell up, resembling elephantiasis (see 
Plate XXIII, fig. 17). 



308 


The ulcers are very chronic, and may persist for many years 
without endangering the life or general health of the patients. 
Judging by the number of natives seen with extensive scars above 
the tibia and with thickening of the bone, it is evident that even 
large ulcers of long standing may suddenly begin to heal up. At 
other times the patients emaciate considerably, become completely 
crippled, and consequently are confined to their houses and die of 
the effects of septic resorption. 

Treatment . Salvarsan has been introduced within the last few 
years for the treatment of tropical ulcer with excellent results. This 
mode of treatment is, however, too expensive and complicated to be 
generally adopted. The thorough removal of the granulation tissue 
with a sharp spoon, followed by careful dressing with a mixture of 
iodoform-boracic acid in the proportion of I to io is efficient. The 
majority of the ulcers yield readily to this treatment, and even large 
sores may heal up within a few months. 

Natives who have been in contact with Europeans for a longer 
period bandage the sores with banana-leaves, and this treatment 
seems to be fairly successful. 

Pathological Histology of Ulcus tropicum 

Opportunity was taken to obtain granulation tissue of io cases 
suffering from Ulcus tropicum in various stages for further histo¬ 
logical examination. The tissue which was preserved in 80 per cent, 
alcohol was embedded in paraffin and sections stained with 
haematoxylin eosin, Breinl’s saffranin-methylene blue, orange-tannin 
(Breinl, 1908), and by Heidenhain’s method, using iodine-potassium 
iodide solution as a mordant. 

The results of histological examination were, on the whole, in 
accordance with those published by Keysselitz and Mayer and by 
Wolbach and Todd. 

The sections of the granulations from the centre of the lesions 
consisted of a loose fibrous mesh-work, the meshes were filled with 
exudate, containing red blood corpuscles, leucocytes and lympho¬ 
cytes, plasma-cells, and a few eosinophile leucocytes were scattered 
here and there. The granulation tissue was richly vascularized, 
many of the arteries showing endarteritis obliterans, and was covered 
by a loose layer of fibrin, which contained erythrocytes in all stages 



309 

of disintegration, lymphocytes, cell detritus and bacilli and spiro- 
chaetes in great numbers. 

The sections from the border of the ulcer showed a marked 
hypertrophy of the epithelial layer, being of many times its normal 
thickness. In between the epithelial cells were roundish or oval 
well-defined spaces of varying size, filled with a loose and 
oedematous fibrous tissue, in which a small number of polymorpho¬ 
nuclear leucocytes were embedded. The epithelial strands between 
these spaces became thinner, the cells showing degenerative changes, 
increasing in size and becoming vacuolated. The nuclei became 
irregular in shape and finally disappeared. In close proximity to 
the ulcers these spaces were densely packed with leucocytes and 
lymphocytes, resembling microscopically small abscesses. 

The superficial epithelial cells from the edge of the sore showed 
degeneration, the cytoplasm was vacuolated, and the nuclei were 
irregular and did not stain evenly. 

In the sections the same microorganisms as described by previous 
observers were found. Fusiform bacilli occurred mostly on the 
surface of the ulcer, and were found in abundance in between large 
numbers of bacteria and cocci. 

The spirochaetes —Spirochaeta schaudinni , Prowazek—to which 
the etiology of ulcus tropicum is generally attributed, were seen in 
small numbers in sections of several of the cases, mostly lying on the 
surface between the other microorganisms or in between the epithelial 
cells adjoining the ulcer. 


CONTRACTING SORE 

A special type of ulcer was encountered, cases of which were 
irregularly distributed throughout the coastal districts of British 
New Guinea. A comparatively greater number of cases was seen in 
the eastern parts, but sporadic cases were discovered in practically 
every district visited. This ulcer differed in its appearance consider¬ 
ably from other sores, and its clinical features were so well marked 
and uniform that it was considered a hitherto undescribed form of 
skin ulceration. It was named 1 Contracting Sore 1 on account of its 
effects. It heals up with formation of dense scar tissue, leading to 
contraction of the joints. 



3io 


The sore appeared in the majority of cases in the neighbourhood 
of joints, for example on the inner or outer surface of the knee 
(fig. 18), on the dorsal surface of the foot, above the tendo achillis, 
or in the neighbourhood of the shoulder joints (fig. 19). Other 
localities, such as around the anus (fig. 20), the abdomen or chest 
or arms were only rarely the seat of the ulcer. On the whole it 
occurred more frequently in young persons, and was often found in 
children. Natives of more mature age only showed the typical scar 
formations. 

The earliest stage was in a boy about 10 years of age. A small 
ulcer of about 3 mm. in diameter was found on the inner side of the 
knee. It was round and possessed irregular burrowing edges; the 
surface was covered by a dry looking, slightly raised, yellowish, 
greasy scab; the surrounding tissue did not show any marked 
inflammation. Underneath this scab, which could easily be 
removed as a whole, was a collection of thick greyish pus accumu¬ 
lated above the granulating surface, which was slightly below the 
level of the skin, of reddish colour, even and of the appearance of 
coarse sandpaper. The granulation tissue was firm and could 
easily be scraped off in strips by means of a sharp spoon. 

The ulcers spread slowly and gradually, at first remaining 
roundish, but after a varying period, becoming irregular in shape, 
and were always covered by the same kind of yellowish-grey scab, 
secreting from the edges an amber-coloured yellowish clear fluid. 
They never spread below the underlying fascia, never attacked the 
bone, and the granulation tissue could always be taken off in whole 
strips, forming a layer of 2 to 5 mm. in depth and exhaling a 
pungent, penetrating odour. 

The ulcer may implicate in some natives a small area only and 
the process may come to a standstill. More often extensive areas 
may be affected, as in fig. 21, where the sore had spread over the 
whole flexor surface of one leg. 

At the time when the ulcerative process is still progressing there 
may be in parts a marked tendency to spontaneous healing (see 
Plate XXIV, fig. 22). The granulating surfaces clear up and are 
replaced by dense and hard scar tissue, possessing a smooth surface, 
and being often extensively intersected by slightly raised strands of 
hard connective tissue. After a varying period, ranging from six 



months to three to four years, the ulcer may completely heal up in the 
way described, giving rise to contraction of the affected parts closely 
resembling at first sight the scar formation due to deep burns (see 
figs. 23, 24). 

This contracture invariably causes further deformities (fig. 25), 
such as complete ankylosis of the joints and wasting of the muscles 
on account of inactivity. 

Histology. From nine cases suffering from different stages of 
1 Contracting Sore * granulation tissue was obtained by scraping the 
sore with a sharp spoon. It was fixed in 80 per cent, alcohol and 
sections made and stained by various methods. The granulation 
tissue from an early ulcer about 3 cm. in diameter showed, on 
microscopic sections, a marked hypertrophy of the epithelial layer 
immediately surrounding the ulcer. These hypertrophied layers of 
epithelial cells were interspersed with irregular spaces filled with a 
loose oedematous connective tissue, the meshes of which contained 
a small number of leucocytes and lymphocytes and a few plasma 
cells; the nearer to the ulcer the larger were these spaces, the thinner 
the epithelial strand separating them and the denser the cell 
infiltration. 

The corium was oedematous and contained a certain amount of 
infiltration of lymphocytes, leucocytes and plasma cells, which 
infiltration was denser around the blood vessels. 

In the central parts of the ulcer the epithelial layer had been 
replaced by granulation tissue, showing here and there a few 
epithelial cells in varying stages of degeneration. The granulation 
tissue itself did not show any distinctive features. 

A number of large cocci and bacteria were seen in sections from 
the surface of the ulcer lying between red blood corpuscles in different 
stages of disintegration and coagulated fibrin. There were neither 
fusiform bacilli nor spirochaetes in the surface layers. In the deepest 
layer of the epithelium, however, in the rete mucosum of the 
malpighian layer especially in the cells lining the spaces referred to, 
a great number of spirochaetes were found, which were either intra¬ 
cellular or were lying in the fine clefts which separated the cells. 
These spirochaetes occurred either singly or sometimes in bunches, 
and stained readily with iron haemotoxylin after iodine-potassium 
iodide had been used as a mordant. The spirochaetes were found 



312 


only in this epithelial layer, and were neither present in the granula¬ 
tion tissue nor in the discharge, of which a small number of smears 
was examined. 

The sections of the granulation tissue of advanced cases did not 
show any especial features. It was richly vascularized, some of the 
vessels showing endarteritis obliterans. There were numerous leuco¬ 
cytes and lymphocytes, a few plasma cells and many phagocytes 
containing small roundish cell inclusions which stained blackish 
with Heidenhain’s iron haematoxylin, and orange when Breinl’s 
method was employed. 

The deepest layers of the granulation tissue consisted of a mesh- 
work of newly-formed dense connective tissue, with numerous fibro¬ 
blasts with a small amount of small-celled infiltration. 

Judging from the histological examination, the ulcer begins with 
a progressive destruction of the epithelial covering of the skin, whilst 
the cells of the deepest layers of the rete mucosum are invaded by 
numbers of spirochaetes. The deep layers of the granulation tissue 
show a tendency to formation of a dense connective tissue. 

On the whole the granulation tissue of these contracting sores 
does not differ microscopically in any essential point from that of the 
ulcus tropicum. The small-celled infiltration is much denser in the 
latter, the surface layers contain a great number of microorganisms, 
fusiform bacilli, bacteria and spirochaetes, whereas in the sections of 
the contracting sore only a very small number of bacilli were found 
in the surface layers, and the spirochaetes were only seen in and 
between the epithelial cells of the rete malpighi. 

The histological examination of the granulation tissue of the 
contracting sore did not reveal any microorganism which could be 
regarded as specific. 

The presence of spirochaetes in numbers in and between the 
layers of the epithelial cells of the malpighian layer is very 
suggestive, but spirochaetes are found in great numbers in many and 
various ulcers of the skin. Moreover, it was impossible to work out 
any detailed morphology of the spirochaete with the material 
collected. 



3^3 

ULCUS INTERDIGITALE DESTRUENS 

A form of ulceration closely resembling ‘ Ulcus interdigitale * as 
described by Castellani and Chalmers (1913) was seen among the 
natives living in the swampy parts of the western New Guinea. 

According to these authors, the ulcer begins as a fissure in 
between the toes, which * Rapidly deepens and enlarges into a large 
oval ulcer with a dull dark red fundus and sodden-looking margins. 
There is practically no discharge whatsoever.’ 

The sores observed resembled, in some respects, those described. 
The ulceration usually started as a fissure between the toes, or more 
often in the sulcus below the toe (PI. XXV, fig. 26). This small fissure 
formed, after a time, into a small painful ulcer, and spread fairly 
rapidly upwards to the toe and downwards to the sole of the foot 
(fig. 27). The ulcer is deep, possessing sharp irregular edges, and 
the granulation tissue, which is covered by an irregular dirty greyish 
scab, has a reddish uneven fundus and discharges a great deal of 
thick yellowish pus. When the ulceration spreads upwards in 
between the toes it causes, on healing, the two adjoining surfaces of 
the toes to grow together. Sometimes the whole toe becomes 
covered with granulation tissue which leads to complete loss of the 
affected toe. Such an example is shown in fig. 28, where the 
ulceration has spread to the sole of the foot, implicating the ball of 
the foot, and had given rise to a deep, irregular granulating sore, 
and two of the toes had been amputated by the ulceration. 

These interdigital sores are very chronic, and show, after a 
shorter or longer period, a tendency to spontaneous healing after 
having caused considerable deformity of the foot. Fig. 29 was 
taken from a case where the fourth and fifth toes had disappeared, 
and where the second and third toes had grown together on their 
adjoining surfaces. 

Cases of this type of ulcer were widely distributed throughout 
the districts of New Guinea west of Orokolo, and affected mostly 
men. The ulcers differ from the ‘ Ulcus interdigitale’ on account of 
their copious discharge and their tendency to lead to the destruction 
of the affected toes, and the name ‘ Ulcus interdigitale destruens * is 
proposed for them. 

Pathological Histology. From two cases suffering from inter- 



3*4 

digital sores, granulation tissue was obtained by scraping the sore 
with a sharp spoon. 

Microscopically the sections did not show any characteristic 
features. The epithelial surrounding of the ulcer was hypertrophic, 
the granulation tissue which was richly vascularised showed a heavy 
leucocytic and lymphocytic infiltration. Numerous bacteria and 
large cocci occurred on the surface, lying in between fibrin cell 
detritus and disintegrating red blood corpuscles. 

GANGOSA 

Gangosa or Rhinopharyngitis mutilans, a disease which causes 
ulceration and destruction of the nose, soft and hard palate, and of 
the skin of the face, is endemic in most of the coastal districts of 
British New Guinea. A full account is being published in this 
journal, describing Cryptococcus mutilans as its etiologic agent 
(sec p. 213). In several far advanced cases the hands and feet were 
the seat of extensive ulceration, the fingers were enlarged, the back 
of the hands swollen and discharged copiously. 

These lesions are in all probability due to the same etiologic 
agent as the lesions of the face. 

SKIN DISEASES 

A great number of different skin diseases are prevalent amongst 
the natives of New Guinea. 

Tinea imbricata (Plate XXVI, fig. 30) was found in every district, 
equally prevalent amongst those natives who had been civilised for 
years, and amongst those who had not come in contact with 
Europeans and had not been away from the district. 

A few cases of Leucoderma were encountered, mostly confined to 
the hands (fig. 31). 

Only two cases of true Albinism were found in the Trobriand 
Islands of the north-east coast of New Guinea—one was in an adult 
man, the other in a baby about three years of age. 

A peculiar form of skin disease was encountered somewhat 
resembling Acne in its clinical appearance, and leading to destruction 
and peculiar scar formation of the affected parts. Sporadic cases of 
this complaint, occurring in men and women, were distributed 



3*5 


throughout British New Guinea. The nose, sometimes the upper 
lip, both cheeks and the forehead, were covered with pustules from 
which sebaceous material could be expressed. The nose became 
enlarged, hyperaemic, and the skin uneven and covered with warty 
excrescences. In one case the upper lip was similarly affected 
(fig. 32). Here and there were small patches of smooth, shiny scar 
tissue, and small roundish depressions with a reddish-looking 
fundus, resembling to some extent the pitting after smallpox 
(fig- 33 )- 

This disease showed a tendency to destroy the affected parts. 
The free border of the alae was uneven and irregular and, in two out 
of eight cases, the soft palate and the uvula had been destroyed and 
replaced by dense scar tissue. 

The skin of the body of the patients was clean and did not show 
any lesions whatsoever. 

As far as histories could be obtained, the complaint seemed very 
chronic, extending over years, and invariably healed spontaneously. 
As far as could be ascertained, the disease made its first appearance 
in the form of pustules on the alae of the nose, whence the lesions 
spread gradually to the cheeks and upper lip. 

An advanced active case is of typical appearance, but when the 
lesions had partly healed the differential diagnosis now and again 
offered difficulties. 

The microscopic examination of the sebaceous material expressed 
from pustules did not give any clue to the etiology. 


CONCLUDING REMARKS 

~~ The material collected during the journey has shown that a great 
many diseases which occur commonly amongst the native popula¬ 
tions in other parts of the tropics are present also in New Guinea. 

Of epidemic diseases only dysentery has made its appearance 
within the last twenty years, and since 1896 outbreaks have been 
recorded from different districts of the Possession. 

A? the primary object of the expedition was the mapping out of 
the distribution of diseases in New Guinea, the short stay did not 
permit a thorough investigation into a number of complaints, 
especially ulcers and skin diseases, many of which could not be 



3*6 

diagnosed and might, on further examination, prove new to 
medicine. 

The two expeditions were made possible by the practical interest 
of the Department of External Affairs, Commonwealth of Australia. 

I take the opportunity of expressing my indebtedness for courtesy 
and assistance received from the Secretary of the Department of 
External Affairs, Mr. Atlee Hunt, C.M.G., from the Lieutenant- 
Governor of Papua, Judge J. H. P. Murray, C.M.G., and his 
personal staff, and from the Government Secretary, Mr. A. M. 
Campbell. 


3*7 


APPENDIX 

LIST OF PAPUAN MOSQUITOS 

*Anopheles (Myzorhynchus) barbirostris , var. bancrofti , Giles. 

Anopheles ( Nyssorhynchus) annulipes , Walker. 

Anopheles (Cellia) punctulata. Ddnitz-Theobald. 

*Armigeres ob turbansf Walker. 

Neosquamomyia breinli , Taylor. 

* Stegomyia fas data, Fabr. 

Stegomyia scutellaris , Walker. 

Stegomyia pseudoscutellaris, Theobald. 

Stegomyia omata , Taylor. 

Stegomyia atra , Taylor. 

Scutomyia notoscripta, Skuse. 

Lepidotomyia lineatus , Taylor. 

Leucomyia australiensis , var. papuensis, Taylor. 

Leucomyia ? albitarsis. Taylor. 

Culicelsa vigilax , Skuse. 

Culex sitiens , var. mi Ini, Taylor. 

Culex fatigans, Wied. 

Pseudotaeniorhynchus conopas . var. giblini , Taylor. 

Chrysoconops brevicellulus , Theobald. 

Taeniorhynchus septempunctata , Theobald. 

Taeniorhynchus uniformis, Theobald. 

Taeniorhynchus papuensis, Taylor. 

Melanoconion papuensis , Taylor. 

Finlaya poicilia , Theobald. 

Uranotaenia nigerrima . Taylor. 

Hodgesia triangulatus , Taylor. 

The bulk of the mosquitos, of which a list has been prepared by Mr. F. H 
Taylor, were collected by Dr. Giblin and the writer at the Lakekamu gold field, 
except those marked with an asterisk, which are previously recorded species. 
(Taylor, 1914). 



3*8 


REFERENCES 

Ashburn and Craig (1907). Observations upon Filaria pbilippensis and its development in 
the Mosquito. Philippine Journal of Science , II, p. 1. 

Bahr (1912). Filariasis and Elephantiasis in Fiji. London. 

Blanchard (1914). Inoculationes experimentales de l’ulcire phagedenique tropical. Bull. 
Soc. Path. Exot., VII, p. 96. 

Breinl (1908). Method of preparing and staining wet films. Annals Trop. Med. and Paras. 
I, p. 470. 

Castellani and Chalmers (1913). Manual of Tropical Medicine. 2nd edition. London. 
Le Dantec (1911). Precis de Pathologie exotique, Paris, 3rd edition. 

Fontoynont et Carougeau (1918). Nodosites juxta-articulaires ; Mycose due au Discomyces 
Carougeaui. Archives de Parasitologic , Vol. XIII, p. 583. 

Fulleborn (1911). Untersuchungen iiber Mikrofilarien in der Siidsee. Archiv f. Scbiffs- 
und T ropenbygiene , XV, p. 368. 

-(1912). Ueber Mikrofilarien des Menschen im deutschen Sudseegebiete und deren 

Turn us nebst Bemerkungen iiber die klinische Manifestation der dortigen Filariasis. 
Arcbiv f. Scbiffs- und Tropenbygiene, XVI, p. 533. 

- (1913). Beitrage zur Morphologic und Differential diagnose der Mikrofilarien. Arcbiv 

f. Scbiffs- und T ropenbygiene, XVII, Beiheft 1. 

Gros (1907). Nodosites juxta-articulaire de Jeanselme chez les indigenes musulman d’Algerie. 
Arcbiv f. Scbiffs- und Tropenbygiene, XI, p. 552. 

Jeanselme (1906). Des nodosites juxta-articulaires observees sur les indigenes de la presqu’ile 
Indo-chinoise. Arcbiv f. Scbiffs- und T ropenbygiene, X, p. 5. 

Joyeux (1913). Contribution A l’etude des nodosites juxta-articulaires. Bull. Soc. Path. 
Exot., VI, p. 711. 

Keysselitz und Mayer (1909). Ueber das Ulcus tropicum. Arch. f. Scbiffs- und Tropenbygiene, 

XIII, p.137. 

Ki nos hit a (1906). Ueber die Verbreitung der Anophelen auf Formosa und deren Beziehungen 
zu den Malariakrankheiten. Arcbiv f. Scbiffs- und Tropenbygiene, X, p. 708. 

Lebouef (1911). Note sur l’existence des nodosites juxta-articulaires de Jeanselme dans 
l’Archipel caledonien. Ann. Hyg. et Mid. Col., p. 549. 

Lynch (1905). Brit. Med. Jour., p. 21. 

MacGregor (1897-1898). Annual Report of Papua, Sanitary, p. XXXIV. 

Neveux (1907). Le Narinde, fibromatose sous-cutanee des Toucouleurs du Boundou (Senegal). 
Rivue Mid. et Hyg. Trop. IV, p. 183. 

Plehn (1914). Die tropischen Hautkranlcheiten. Handbucb der Tropenkrankbeiten, Leipzig, 
Vol. II, p. 171. 

Prowazek (1907). Vergleichende Spirochaetenuntersuchungen. Arb. a. d. Kaiserl. Gesund - 
beitsamt, XXVI, p. 23. 

Steiner (1904). Ueber multiple subkutane harte fibrose Geschwiilste bei den Malayen. 
Arcbiv f. Scbiffs- und Tropenbygiene, VIII, p. 156. 

Taylor, F. H. (1914). Culicidae from Papua. Trans. Ent. Soc. London, Part I, pp. 185-205. 
Pis. XVII, XVIII. 

Taylor, F. H. (1914). Contributions to a knowledge of Australian Culicidae, No. 1. Proc. 
Linn. Soc. N. S. Wales, Part 3, pp. 454-468. Pis. XXXIV, XXXVII. 

Thorpe (1896). Brit. Med. Journ., II, p. 922. 

Wolbach and Todd (1912). A study of chronic ulcers, Ulcus tropicum, from the Gambia. 
Journ. Med. Research, XXII, p. 27. 




320 




EXPLANATION OF PLATES 



Plate XIX 

Fig. 

i. 

Elephantiasis of scrotum. 

Fig. 

2. 

Elephantiasis of vulva. 

Fig. 

3- 

Leprosy; advanced case. 

Fig. 

4- 

Juxta-articular nodules. 



Annals Trop. Med. o Parasitol ., Vol. IX 


PLATE XIX 



Fig. i 


Fig. 3 



F 'c- 2 Fig. 4 



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322 


Plate XX 

Fig 5- Juxta-articular nodules. 

Fig. 6. Yaws. General eruption. Secondary stage. 

Fig. 7. Yaws. General eruption. The same case as fig. 6. 

Fig. 8. Yaws. Note destruction of nose. 




Fig. 5 Fig. 6 



Fig. 7 Fig. 8 


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3 2 4 


Plate XXI 

Figs. 9-12. Disease characterised by osteitis, periostitis, arthritis, 
and formation of sores. 

Fig. 9. Late stage, resembling Madura foot. 

Fig. 10. Swelling of bone affecting the tibia. 

Fig. II. Swelling of bone affecting the diaphysis of radius. 

Fig. 12. Mutilation of hand after resorption of metacarpal 
bones. 



Annals Trop. Med. & Parasitol ., Vol. IX 


PLATE XXI 



Fig. ii 


Fig. 12 


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326 


Plate XXII 

Figs. 13-14. Disease characterised by osteitis, periostitis, arthritis, 
and formation of sores. 

Fig. 13. Metatarsus of large toe disappeared, the toe in 
hyper-extension and abduction. 

Fig. 14. Late stage of disease, showing irregular scar 
formation. 

Figs. 15-16. Ulcus tropicum. 





Annals Prop. Med. & ParasitolVol. IX 


PLATE XXII 


Fig. 14 


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Plate XXIII 


Fig. 17. Ulcus tropicum. 
Figs. 18-21. Contracting sore. 



Annals Trop. Med. G? Parasitol., Vol. IX 


PLATE XXIII 




Fie. 19 


Fig. 21 


Fig. 20 


Fig. 17 


Fig. 18 


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33 ° 


Figs. 22-25. 


Plate XXIV 


Contracting sore. 



Annals Trop. Med. £*? ParasitolVol. IX 


PLATE XXIT 



% 

x 


•v 


Fig. 24 


Fig. is, 


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332 


Plate XXV 


Figs. 26-29. Ulcus interdigitale destruens. 



Annuls 'Trot>. Med. ParasitolVol. I .Y 


PLATE XXV 



Fig. 27 Fie. 29 


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334 


Plate XXVI 

Fig. 30. Tinea imbricata. 

Fig. 31. Leucoderma. 

Figs. 32-33. Skin disease resembling Acne. 


Annals Tro[>. Med. ZA Paras itol., Vol. IX 


PLATE XXVI 



Fic. 31 Fig. 33 


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335 


INSECT FLAGELLATES AND THE 
EVOLUTION OF DISEASE, WITH 
REMARKS ON THE IMPORTANCE 
OF COMPARATIVE METHODS IN 
THE STUDY OF PROTOZOOLOGY 


BY 

H. B. FANTHAM, M.A. Cantab., D.Sc. Lend. 

LECTURER ON PARASITOLOGY, LIVERPOOL SCHOOL OF TROPICAL MEDICINE 


{Received tor publication l May , 1915) 

CONTENTS 

PAGE 


I —Introduction . 335 

II —The Significance of the Herpetomonad Stage of Ltisbmania . 337 

III — The Existence of the Herpetomonad Stage of Leisbmania and Allied 

Parasites in Man . 339 

IV — The Presence of Herpetomonads in Nature in other Vertebrates ... 339 

V— The Occurrence of Herpetomonads in Plants . 341 

VI —Experiments with Insect Flagellates belonging to the Genera Hcrpetomonai 

and Critbidia . 342 

VII — Some Inferences . 344 

VIII — Summary . 346 

References . 347 


I. INTRODUCTION 

The parasitic theory of disease has been greatly extended and 
consolidated during the last generation. In the domain of 
bacteriology it has perhaps become best known, at any rate to the 
general public. Vast strides have been made during the latter half 
of the period in our knowledge of animal parasites, especially of the 
inter-relationship of Protozoa and arthropods in the propagation of 
disease. There is also a re-awakening as to the importance of 
mycology, or the study of fungoid organisms as pathogenic agents 
of economic importance. 

In the domain of protozoology, with which the present paper is 
concerned, the progress, although rapid, has not been so great as it 
might have been. Some of the reasons for this are that the work 



is in the main in the hands of medical men whose knowledge of 
zoology is generally of the slightest and their acquaintance with the 
comparative aspect of the subject just as limited, while the few 
zoologists engaged have in some cases drifted into a narrow outlook 
and become lost in academic terminologies and discussions. The 
result is that though many important discoveries have been made, 
they have not reached full fruition, chiefly on account of lack of 
application of the comparative method. As an example, the parasite 
of kala-azar may be considered. The causal organism, Leishmania 
donovani , was described in 1903 by Leishman and by Donovan, 
while Rogers in 1904 cultivated it and discovered the herpetomonad 
stage. The latter discovery, that of Rogers, was of fundamental 
importance, but its application is, even to-day, only realised in 
varying degrees by two or three British and French workers. 
However, the output of literature on kala-azar and other 
leishmaniases is simply enormous, though the progress resulting 
therefrom is very slight, because of the overburdening amount of 
controversy obscuring the issue. When Rogers in 1905 suggested 
the bed-bug as the probable transmitter of kala-azar in India, and 
Patton began to work at the same, the results obtained were largely 
buried in criticism, much of it speculative and irrelevant. We read 
in 1912 such statements as the following : ‘ It is quite clear that the 
gut of the bug when it contains blood acts as a culture-tube 
in which various organisms can live and develop. . . . Hence the 
fact that the leishmania of kala-azar and oriental sore become 
flagellates in the bug’s gut is in itself no proof whatever that 
these insects are the transmitter’s of these diseases.* It is to be 
regretted that those putting forward or supporting the latter 
remarks did not more carefully consider the suggestions and 
inferences to be drawn from cultural methods, and proceed forthwith 
to experiment directly with the herpetomonads found in insects, 
especially after Patton (1907) had given the first account of the 
complete life-cycle of such a herpetomonad, and Darling (1910) had 
suggested that a case of oriental sore in Panama had resulted from 
' an inoculation with an invertebrate gut flagellate (Crithidia ?).* 
Again, we had the flood-gates of destructive criticism opened when 
Basile (1910-11) suggested that Leishmania infantum was 
transmitted by fleas, though it does not seem to have occurred to 


337 


any of the critics to begin experiments on the effects of flagellates of 
fleas when introduced into the vertebrates they infest. However, it 
is unnecessary to labour the point further, it is merely symptomatic 
of the age, pointing to the partial failure of the modern educational 
system, in that it tends to the development of trifling academic 
debate, nearly always useless and sometimes even harmful, and to a 
fostering of the partisan spirit, wherein a broad comparative outlook 
is seldom attained. Legitimate criticism is always welcome, but 
when it degenerates into mere retort and negation it is subversive 
of progress. 

There is yet another branch of the study of pathogenic organisms 
which has not yet received attention, namely, the biological evolution 
of such forms as disease producers, and a consideration of ‘ disease 
in the making.* For this study a knowledge of comparative 
morphology and of the inferences to be made therefrom is absolutely 
essential, and any one who aspires to become a parasitologist must 
be prepared to undertake such a study. 

II. THE SIGNIFICANCE OF THE HERPETOMONAD STAGE OF 

LEISHMANIA 

When Rogers in 1904-05 cultivated the Leishman-Donovan body, 
obtaining a herpetomonad flagellate therefrom, and suggested that 
the genus Leishmania should give place to Herpetomonas , a great 
step forward might and should have been made in the application 
of the idea, especially as Christophers (1904) in L. donovani , and 
Mesnil and colleagues (1904) in L. tropica , had shown that a 
structure, now known to be the root of the flagellum, could be found 
in some of the parasites. ** Patton (1908) also placed Leishmania 
donovani in the genus Herpetomonas. It was also pointed 
out that the natural herpetomonads of insects might be mistaken 
for developing Leishmania. However, these advances met 
with the inevitable destructive criticism, for as recently as 1912 
we read: ‘ The genus Herpetomonas includes flagellates which 
have only one, and that an invertebrate host. The fact that 
the parasite of kala-azar has two hosts and can live and 
multiply in the organs of a vertebrate shows it to be profoundly 

• Novy (1909) »howed that a rhizoplast occurred also in /.. infantum. (Bull. Soc. Path. 

Exol., II, pp. 385-387.) 



338 


different from the true Herpeiomonas , which lives only in the 
invertebrate. This fact alone would be sufficient to establish the 

genus Leishmania .When a Herpetomonas becomes so 

changed that it has acquired the power of living and developing as 
an intracellular parasite in the body of a warm-blooded animal, 
and giving rise to such a serious disease as kala-azar, 
we are justified in concluding that it has passed out of the 
genus Herpetomonas , from which it originated, into the genus 
Letshmania* The pity is that the author, before penning 
these remarks, did not himself endeavour to determine experi¬ 
mentally whether Herpetomonas might not acquire the power 
of living and developing in a vertebrate, and that the record of the 
remarkable observations of Dutton and Todd, published in 1903, 
on the presence of herpetomonads in the blood of Gambian house 
mice was unknown to him, as to many others. I deeply regret the 
necessity for making these remarks; there is nothing personal in 
them, but the facts are there and cannot be gainsaid. 

The successful cultivation of L. tropica and L. infantum again 
did not advance matters, but discussion arose as to the number of 
valid species of Leishmania y and numerous attempts were made to 
inoculate the viruses into laboratory mammals. Although expedi¬ 
tions were organised for the study of Leishmania in endemic areas, 
it never seems to have occurred to those working thereon to try 
to introduce herpetomonad flagellates of insects into vertebrates. 
Apparently, vertebrates, being thought much more powerful and 
important animals than invertebrates, must perforce be primarily 
considered and experimentation undertaken with them viewed 
as the principal hosts. Unfortunately, human suffering has paid 
dearly for this one-sided view, which is only equalled by the 
ridiculous arguments, that used to occupy the attention of amateur 
parasitologists and waste the time of learned societies, as to which 
was the primary or principal host of a digenetic parasite, was it the 
vertebrate or the invertebrate host ? The answer is that each host 
has its own particular importance which must be carefully considered. 
In the case of the kala-azar parasite the pathogenic organism 
certainly seems to have arisen from a herpetomonad parasite of an 
invertebrate (probably an insect), and the flagellate has acquired the 
capacity to live within a vertebrate and so become digenetic. 




339 


Again, the intimate connection of natural canine leishmaniasis 
and of infantile kala-azar has been explained away by some. In 
other words, every step forward has been obscured by irrelevant 
criticism, and what should have been obvious inferences were either 
ignored or never given the opportunity of being put to the practical 
test. The comparative aspect was overlooked. The occurrence of 
natural herpetomonads in invertebrates must not only be acknow¬ 
ledged, but it must be allowed that they may become pathogenic 
when introduced into vertebrates. 

III. THE EXISTENCE OF THE HERPETOMONAD STAGE OF 

LEISHMANIA AND ALLIED PARASITES IN MAN 

It is now known that Escomel in Peru in 1911 found the herpeto- 
monad stage of Leishmania tropica in man, but seeming somewhat 
disturbed by his finding did not publish it at the time, nor realise 
its importance and significance. Monge, also in Peru, records 
(1914) the presence of the herpetomonad stage in dermo-mucosal 
leishmaniasis, and La Cava (1912) in L. tropica in Italy. Splendore 
(1912) saw elongating forms and a few flagellate stages in cases in 
Brazil. 

Even before these observations, Darling (1906, 1909) in Panama 
had recorded a disease resembling kala-azar, and described the 
causal parasite therein as His toplasma capsulatum. He stated that 
flagellate stages occurred in the lungs and spleen. As might be 
expected, the mention of the occurrence of flagellate stages passed 
almost unnoticed. In 1912 Rocha-Lima, working in Hamburg, 
stated that in his opinion Histoplasma capsulatum was a 
Blastomycete. Whether this be so or not, surprisingly little 
attention was paid to Darling’s researches. 

Again, in 1913 Franchini described a herpetomonad parasite 
from man, and called it Haemocystozoon brasiliertse. 

IV. THE PRESENCE OF HERPETOMONADS IN NATURE IN 

OTHER VERTEBRATES 

Dutton and Todd in 1903 recorded the presence of herpetomonad 
flagellates in the blood of mice in the Gambia. Unfortunately, these 
observations, first made in 1902, seem to have escaped general 
notice. They have since been interpreted otherwise by the surviving 



34 ° 


author. Three Gambian house mice out of fourteen were found to 
be infected with flagellate herpetomonads, of which a clear description 
is given, together with dimensions. The infection was scanty, and 
the parasites were only seen in fresh preparations. The organisms 
were definitely compared with Herpetomonas ( Leptomonas) 
biitschlii recorded from the Nematode worm, Trilobus gracilis . 

A diversion is necessary here. The parasite, Herpetomonas 
( Leptomonas) biitschlii , briefly described by Biitschli in 1878 and 
named by Saville Kent in 1880, emphatically needs re-investigation. 
According to Biitschli, the organism was spindle-shaped or almost 
rod-shaped, bearing a long flagellum and possessing a contractile 
vacuole—a feature not present in flagellates latterly known as 
‘ leptomonads.* No nucleus was observed by him. A darker, 
refractile granular mass behind the contractile vacuole may have been 
the blepharoplast, or even the nucleus. Saville Kent appears to have 
considered it to be the nucleus, and he incorrectly reproduced it as a 
vesicle in his ‘ Manual of the Infusoria.* The developmental cycle 
was unknown to Biitschli, and still remains so. The organism has 
not been studied by modern methods of technique, probably because 
of the difficulty of obtaining the host. It seems rather remarkable 
that a parasite which is somewhat shrouded in mystery should be 
made the type species of a genus, Leptomonas , which some workers 
would like to see supersede the well-known genus Herpetomonas . 
Moreover, Biitschli in 1884, in his account of the Mastigophora in 
Bronn*s ‘Tierreich,* considered the two generic names to be 
synonymous and retained the name Herpetomonas. The account of 
H. muscae domesticae by Prowazek having been shown by many 
competent investigators to be inaccurate—the so-called bi-flagellate 
characteristic being due to division—there is no need for the 
multiplication of genera and controversy on the same. 

To return to the natural occurrence of herpetomonads in verte¬ 
brates. That these flagellates may occur naturally in mice is 
undoubted, as they have been seen and described by other workers 
in addition to Dutton and Todd. In May, 1909, while working in 
the Quick Laboratory, Cambridge, a Herpetomonas was found in 
life in the fresh peripheral blood of a mouse. It was seen almost 
simultaneously and independently by Prof. Nuttall, Dr. Porter and 
myself, but the infection was slight. The parasite has been seen 


since, but again the infection was scanty. These observations have 
just been published by Fantham and Porter. The parasite is very 
probably Herpetomonas pattoni (Swingle), first seen in rat-fleas, as 
some of these ectoparasites were breeding in the rat cages in the 
animal house in which the mice were kept. 

The so-called herpetomonad stages of trypanosomes are probably 
natural herpetomonads of vertebrates co-existing with trypanosome 
infections. This subject is discussed on page 345. 

V. THE OCCURRENCE OF HERPETOMONADS IN PLANTS 

In 1909 Lafont announced the discovery in Mauritius of a 
Herpetomonas —or Leptomonas , as he called it—in the latex of a 
species of Euphorbia or spurge. Confirmation followed rapidly, and 
the flagellate (H . davidi) is now known in various species of 
Euphorbia in neighbouring islands, in India, Africa, Portugal, and 
other places (see Franca, 1914). In some of these places Hemiptera 
were found crawling over the plants, and herpetomonads were also 
found in the gut of the Hemiptera. Some of the parasitised plants 
were diseased, and the malady has been termed 4 flagellosis.* 

The flagellate parasite of the spurge plants is probably an insect 
flagellate, coming from the Hemiptera infesting the plants. The 
herpetomonad has probably secondarily invaded the plant tissue, 
having adapted itself so that it is now able to live in the latex of the 
plants. Encysted or post-flagellate stages of H. davidi have been 
obtained in cultures. 

Nearly three years ago I was informed, by a competent 
authority, that a number of Euphorbia containing herpetomonads 
grew outside a certain hospital situated in an area in which kala-azar 
was endemic, and in which kala-azar patients were being treated. 
The shrubs were infested by insects. It seems remarkable that no 
attempt was made to trace a possible connection between the plant 
herpetomonad and kala-azar, doubtless such a possibility was 
considered too remote. Remarks of mine regarding a possible 
connection were 'received by my informant with polite incredulity, 
which is not surprising, since the wisdom of lecturing on 
Herpetomonas and Crithidia to students of tropical medicine has 
been questioned more than once. 



34 2 

VI. EXPERIMENTS WITH INSECT FLAGELLATES BELONGING 
TO THE GENERA HERPETOMONAS AND CRITHID1A 

Thanks to the remarkable series of experiments recently carried 
out by Laveran and Franchini in Paris, and by Fantham and Porter 
in England, we now know that a number of species of Herpetomonas 
and Crithidia , naturally occurring in insects, may be successfully 
inoculated into or fed to mammals, especially rats and mice. The 
flagellates live and multiply in the vertebrates and become pathogenic 
thereto, the symptoms chiefly resembling those of kala-azar. It 
must not be supposed, however, that success is attained in every 
experiment, though a large proportion are positive, especially if the 
vertebrate hosts are young. (Compare the prevalence of leishmaniasis 
in young people (children) in the Mediterranean region.) 

In the insect host the Herpetomonas or Crithidia is a natural or 
specific parasite, and its effects on its host are not marked. The 
life-cycle of the organism in the invertebrate consists of a 
leishmaniform pre-flagellate stage, gradually growing into a 
flagellate form and followed by a resistant post-flagellate, 
leishmaniform stage adapted for extra-corporeal life and for 
transmission to a new host. When such a parasite finds its way, 
either by inoculation or by feeding, into a susceptible vertebrate, it 
can assume again leishmaniform or flagellate facies. The mode of 
infection of the vertebrate in nature seems to be contaminative, either 
by its food or through an already existing abrasion or puncture on 
the surface of its body. Cases in which the flagellate-infected 
insects have been allowed to suck the blood of vertebrates have proved 
negative. Further, Fantham and Porter have brought forward 
experimental evidence which shows that post-flagellate forms of the 
parasites are best adapted to begin life in a vertebrate host. 

In certain cases leishmaniases may be derived from reservoir 
animals, such as man and various other warm- and cold-blooded 
vertebrates (see page 344), in other cases they may arise de novo. 

Laveran and Franchini have successfully introduced into 
mammals Herpetomonas ctcnocephali from the gut of the dog-flea, 
H . pat tout from the gut of the rat-flea, Crithidia jasciculata from the 
gut of Anopheles maculipennis, and C. melophagia from the gut of 
the sheep-ked. 

Experiments have also been made with H. muscae domesticae. 
The mammals used were chiefly rats and mice. 


3+3 


Fantham and Porter (1914) went further, and showed that 
flagellates occurring in insects unassociated with the experimental 
vertebrate may be introduced therein. Their conclusions were as 
follows: — 

1 1. Insect flagellates, e.g., Hcrpetomonas jacutum (Leger) 
from Nefa einerea , and Herpeiomonas ctenocephali 
(Fantham), parasitic in the dog-flea, Ctenocephalus cants , 
can live inside certain vertebrates (e.g., mouse and dog, 
respectively) and can multiply therein. This we have 
shown experimentally. 

‘ 2. If such flagellates be inoculated intraperitoneally, or are 
fed by the mouth in food, the flagellates can find their way 
into the blood stream and internal organs (e.g., liver, 
spleen, bone-marrow) of the vertebrate host. 

‘ 3. The insect flagellates are pathogenic to the vertebrates 
experimented upon, producing symptoms like those of 
leishmaniasis (kala-azar). 

‘ 4. The oval post-flagellate forms appear to be more capable 
of developing in vertebrate hosts than are other stages of 
tjie herpetomonad parasite of the insect. 

‘5. It may be expected that the various leishmaniases, 
occurring in different parts of the world, will prove to 
be insect-borne herpetomoniases.* (According to some, 
leptomoniases would probably be considered correct, but 
see page 340.) 

In a later paper further experiments were recorded by Fantham 
and Porter, and the researches were also extended to cold-blooded 
vertebrates. The conclusions were : — 

* 1. Herpetomoniasis can be induced in various warm- and cold¬ 

blooded vertebrates when the latter are inoculated or fed 
with herpetomonads occurring in the digestive tracts of 
various insects. The infection produced and the protozoal 
parasites found in the vertebrates resemble those of human 
and canine leishmaniases. 

* 2. An infection can also be induced in certain vertebrates 

when they are fed or inoculated with Crithidia gtrridis , 
and both flagellate and non-flagellate stages occur therein, 
but no transition to a trypanosome was found. 



3+4 


‘ 3. The following Flagellata have proved pathogenic to warm¬ 
blooded mammals when the latter have been fed, or 
inoculated subcutaneously or intraperiloneally with them— 
Herpetomonas'jaculum , H. stratiomyiae , H. pediculi and 
Crithidia gerridis . The hosts used were mice of various 
ages. That H. ctcnoccphali can infect dogs has already 
been shown by us. 

‘4. Herpetomonas jaculum and Crithidia gerridis have also 
been successfully fed or inoculated into cold-blooded 
hosts, namely, fishes ( Gasterosteus aculeatus ), frogs, 
toads, lizards ( Lacerta vivipard) and grass snakes 
(Tropidonotus natrix). 

‘5. As we have previously stated, we believe that leishmaniases 
are arthropod-borne herpetomoniases, and that these 
maladies have been evolved from flagellates of inverte¬ 
brates (especially herpetomonads of insects) which have 
been able to adapt themselves to life in vertebrates. 

4 6. In areas where leishmaniases are endemic, an examination 
should be made of all insects and other invertebrates likely 
to come into contact with men or dogs or rats and mice, 
in order to ascertain if these invertebrates harbour herpeto¬ 
monads. Preventive measures should be directed against 
such invertebrates, especially arthropods. Further, it is 
likely that certain vertebrates, such as reptiles and 
amphibia (especially those that are insectivorous), may 
serve as reservoirs for leishmaniases or, as they should 
preferably be termed, herpetomoniases. From such 
reservoirs the herpetomonads may reach man by the 
agency of ectoparasites or flies, especially such as are 
sanguivorous.* 

VII. SOME INFERENCES 

The immediate and important inference to be drawn from the 
experiments recorded in the previous section is that in them we sec 
4 leishmaniasis in the making,’ as was pointed out by Fantham 
and Porter in December, 1913, in these Annals. Also canine 
leishmaniasis is probably herpetomoniasis due to H. ctenocefhal'i 
in dogs. 

A further imperative inference is that probably only one species 



3+5 


of Herpetomonas is concerned in adapting itself to life in vertebrates 
in different parts of the world. This species is known under various 
names, such as H . pattoni , H . ctenocephali , H. pediculi , and also as 
H . donovani , //. infantum , //. tropica . These are probably merely 
physiological races of a herpetomonad which is very like //. jaculum , 
briefly described by L6ger in 1902 from the gut of the Hemipteran, 
cinerea. This herpetomonad under different conditions of 
environment produces pathogenic effects in very varying degrees in 
different vertebrates, from zero as in Dutton and Todd’s mice to 
high mortality as in Indian kala-azar, and probably zero again in 
cold-blooded hosts. It is also a flagellate which can probably live 
in invertebrates not already recorded as being infected. 

At various times statements have been made that herpetomonad 
stages of trypanosomes occur in cultures. The invertebrate life- 
cycles of all the trypanosomes, which so far have been well investi¬ 
gated, lend no support to the presence of a herpetomonad stage 
therein. It is highly probable that the so-called herpetomonad 
stages of trypanosomes were really cultures of scanty herpetomonad 
infections co-existing with trypanosome infections. -It is known 
that such invertebrates as rat-fleas may be infected with both 
herpetomonads and trypanosomes at the same time. The different 
discussions in the past, often acrimonious, as to what form of 
flagellate occurring in an invertebrate constituted part of the life¬ 
cycle of a trypanosome or otherwise, are thus to be deplored. 

Before concluding, attention may be drawn to some remarkable 
and far-seeing suggestions regarding kala-azar published by 
Rogers in 1905. He wrote that 4 the stomach of some blood¬ 
sucking insect is the most likely place in which to find 
the natural development of the extra-corporeal stage of the 
parasite, and that some such insect is the most likely carrier 

of the infection.But as to the most likely kind of insects 

to carry the infection we are on more uncertain ground, for 
experiment alone can determine this. Nevertheless, it is worth while 
to discuss which are the most probable kinds in order that 
precautions may be taken against them without waiting for absolute 
proof to be obtained.’ Especial notice is drawn to the last sentence, 
published ten years ago, but not yet acted upon fully from the point 
of view of preventive measures. However, exception must be made 
for the excellent researches and preventive measures undertaken by 




346 


Dodds Price, at the suggestion of and latterly in collaboration with 
Rogers, in the Assam tea gardens. He has reduced the mortality 
due to kala-azar enormously by segregating the infected, by moving 
coolie lines about 300 yards from older infected ones, and by having 
new coolie lines placed on clean sites. 

It has remained for workers in laboratories in temperate zones, 
away from leishmaniasis material, again to point the way to 
preventive measures and to indicate the origin of the disease. 

VIII. SUMMARY 

The significance of the herpetomonad stage of Leishmania y of 
the recent announcements that such stages occur in man, and of the 
presence of natural herpetomonads in other vertebrates (for example, 
mice), are discussed. It also recalled that insect herpetomonads can 
invade and live in plant-tissues. 

The experiments on the introduction into different vertebrates of 
various species of Herpetomonas and Crithidia parasitic in insects 
by Laveran and Franchini, using mammals, and by Fantham 
and Porter, using both warm- and cold-blooded vertebrates, are 
summarised and discussed. 

It is inferred that the various leishmaniases are due to a 
herpetomonad of invertebrates which, under different conditions of 
environment, produces pathogenic effects in very varying degrees in 
different vertebrates, from zero, as in the mice described by Dutton 
and Todd in 1903, to high mortality as in Indian kala-azar, and 
probably zero again in cold-blooded hosts. It is also a flagellate 
which can probably live in invertebrates not already recorded as 
being infected. A human reservoir of leishmaniasis may occur in 
some places, while warm- and cold-blooded vertebrates may also 
function as the same. 

It is highly probable that the so-called cultural herpetomonad 
stages of trypanosomes were really cultures of scanty herpetomonad 
infections co-existing with trypanosome infections. 

It is recalled that Rogers in 1905 published that ‘it is worth 
while to discuss which are the most probable kinds [of insect 
transmitters of kala-azar] in order that precautions may be taken 
against them without waiting for absolute proof to be obtained.' 
Although these remarks were published ten years ago, little has been 
done directly in the way of the preventive measures suggested. A 
notable exception, however, is the work of Dodds Price in Assam. 



347 


Addendum. —June 28, 1915. 

Since writing this paper my attention has been drawn to a recent 
note by Archibald, who writes; ‘ Epidemiological and experimental 
evidence does not support the theory that kala-azar in the Sudan is 
transmitted by a biting insect. A more probable source of infection 
appears to be some intermediate host, whose habitat is in water/ 
{Kept, of Advisory Committee for Trop. Dis. Resch. Fund for 1914, 
p. 116 —published in April, 1915.) In this connection, see my 
remarks on the contaminative method of infection in paragraph 2 
on page 342. 


REFERENCES 


Further reference* will be found at the ends of some of the memoirs cited. 


Basils, C. (1910). Sulla Leishmaniosi del cane e sulP ospite intermedio del Kala-Azar infantile. 
Rendiconti R. Accad. dei Lincei , (ser. V.), XIX, ii, pp. 523-527 

-(1911)* Sulla Leishmaniosi e sul suo modo di trasmissione. Rendiconti R. Accad. dei 

Lincei , (ser. V.), XX, i, pp. 278, 479, 955. Sec also p. 50. 

Botschli, O. (1878). Beitrage zur Kenntniss der Flagellaten und einiger verwandten 
Organismen. I. Zeitscbr. f. toissen. Zoologie, XXX, pp. 205-281. 5 plates. 

Christophers, S. R. (1904). On a parasite found in persons suffering from Enlargement 
of the Spleen in India—second report. Sci. Mem. Govt. India , No. 11. 

Darling, S. T. (1909). The Morphology of the Parasite ( Histoplasma capsulatum) and the 
Lesions of Histoplasmosis, a fatal disease of Tropical America. Journ. Exptl. Med., 
XI, pp. 5 * 5 - 53 *- splato. 

*- (1910). Autochthonous Oriental sore in Panama. Trans. Soc. Trop. Med. 

® Hyg > rv, pp. 60-63. 

Dutton, J. E., and Todd, J. L. (1903). ‘ Flagcllata in the Blood of a Mouse,' being part of 

Sect. VII of the First Report of the Trypanosomiasis Expedition to Senegambia (i 9 ° 2 )- 
Liverpool Scb. Trop. Med., Memoir XI, pp. 56-57. 

Escomel, E. (1914). Leishmania flagelada en el Peru. La Cronica Medica. [Lima], XXXI, 
pp. 224-227. See also (1913) Bull. Soc. Path. Exot ., VI, p. 237. 

Fantham, H. B. (1912). Some Insect Flagellates and the Problem of the Transmission of 
Leishmania. Brit. Med. Journ., Nov. 2, 1912, pp. 1196-1197. 

Fantham, H. B., and Porter, A. (1913). Herpetomonas stratiomyiae , n.sp., a flagellate 
parasite of the flies, Stratiomyia chameleon and 5 . potamida, with remarks on the biology of 
the hosts. Annals Trop. Med. and Parasitol., VII, pp. 609-620. 1 plate. 

-(1914-15). Some Insect Flagellates introduced into Vertebrates. Proc.Camb. 

Pbilosopb. Soc., XVIII, pp. 39-50. 1 plate. Also Cambridge Univ. Reporter , XI.V, 

p. 323 (Dec. 1, 1914). 

-(1915). Further Experimental Researches on Insect Flagellates introduced into 

Vertebrates. Proc. Camb. Pbilosopb. Soc., XVIII, pp. 137-148. Also Cambridge Univ. 
Reporter , XLV, p. 931 (May, 25, 1915). 

-(1915). On the Natural occurrence of Herpetomonads (Leptomonads) in Mice. 

Parasitology, VIII, pp. 128-132. 



3+8 


Franca, C. (1914) Fa Flagellose des Euphorbes. Arch. f. Protistenkunde , XXXIV, 
pp. 108-132. 1 plate. 

Francium, (». (1913;. L’n nouveau protozoairc parasite de l’homme provenant du Brasil. 
Bull. Soc. Path. Fxot VI, pp. 156-158. See alio pp. 333-336. 

La Cava, F. (1912). De la Leishmaniose dci muqueuses et de la premiere dFcouverte de la 
Lcisbmania tropica flagellcc dans lc corps humain. Bull. Soc. Path. Exot ., V, pp. 808-812. 

Lav Fran, A., and Franchini, G. (1914). Infections de mammiferes par des Aagcllts d’invcrtF- 
bres. Bull. Soc. Path. Exot., VII, pp. 605-612. 

Mesmi., F., Nicolle, M., and Remlinger, P. (1904). Sur lc protozoaire du bouton d’Alep. 
C.R. Soc. Biol., LVII, pp. 167-169. 

Mover., l\ (1914). La Leishmaniasis del dermis en el Peru. Espundia, Uta, Juccuya, Qcepo, 
Tiacc-arana. La Cronica Medica. [Lima], XXXI, pp. 231, 251, 288, 385. 

Patton, W. S. (1907;. The Development of the Leishman-Donovan parasite in Cimex 
rotundat us. Second report. Set. Mem. Govt. India, No. 31. Sec also No. 53. 

-(11)07). Preliminary Note on the Life-cycle of a Species of Herpetomonas found in 

Culex pipiens. Brit. Med. Journ ., July 13, 1907, pp. 78-80. 

— (1908). Inoculation of dogs with the parasite of Kala azar ( Herpetomonas [Leisbmania] 
donmant), with some remarks on the genus Herpetomonas. Parasitology, I, pp. 311-313. 

Porter, A. (1914). The Morphology and Biology of Herpetomonas patellae, n.sp., parasitic 
in the Limpet, Patella vulgata , together with remarks on the pathogenic significance 
of certain Flagellates found in Invertebrates. Parasitology, VII, pp. 322-329. 

Price, J. Dodds, and Rogers, Leonard (1914). The uniform success of segregation measures 
in eradicating Kala-azar from Assam tea gardens. Its bearing on the probable mode 
of infection. Brit. Med. Journ., Feb. 7, 1914, pp. 285-289. 

Rocha-Lima, H. da (1912). Histoplasmosis und epizootische Lymphangitis. Arch. j. 
Scbiffs- it. Tropenbvgiene. XVI, Bcibeft 1, pp. 79-85. 

Rogers, L. (1905). The Conditions affecting the Development of Flagellated Organisms 
from Leishman bodies and their bearing on the probable mode of Infection. Lancet , 
June 3, 1905, pp. 1484-1487. 

Spi.endore, A. (1912). Lcishmaniosi con localizzazione nelle cavita mucose (nuova forma 
clinica). Bull. Soc. Patb. Exot., V, pp. 411-438. 2 plates. 

Wi n yon, C. M. (1912). Experiments on the behaviour of Lcishmania and Allied Flagellates 
in Bugs and Fleas, with some Remarks on Previous Work. Journ. Lond. Scb. Trap. 
Med., II, pp. 13-26. 



1 


Volume IX 


July. 1915 


No. 3 


ANNALS 

OF 

TROPICAL MEDICINE AND 
PARASITOLOGY 


ISSUED BY 

THE LIVERPOOL SCHOOL OF TROPICAL MEDICINE 


Edited by 

Professor J. W. W. STEPHENS, M.D. Cantab., D.P.H. 

Professor R. NEWSTEAD, M.Sc., J.P., F.R.S., A.L.S., F.E.S., Hon. F.R.H.S. 
Professor WARRINGTON YORKE, M.D. 

AND 

Professor Sir RONALD ROSS, K.C.B., F.R.S., M.D., F.R.C.S., 

Major I.M.S. (Ret.) 

Editorial Secretary 
Dr. H. B. FANTHAM, 

School of Tropical Medicine , 

The University , 

Liverpool. 



C. Tinting Co., Ltd. 

Printers to the University Press of Liverpool 
53 Victoria Street 



3.49 


NOTES ON THE BIONOMICS OF 
GLOSSINA PALPALIS IN SIERRA 
LEONE, WITH SPECIAL REFERENCE 
TO ITS PUPAL HABITATS 

[Being the First Report of the Thirty-second. Expedition of the 
Liverpool School of Tropical Medicine, 1914-1915] 

BY 

WARRINGTON YORKE 

AND 

B. BLACKLOCK 

(Received for publication 21 April, 1915) 

Plates XXVII-XXXIII and Map 

INTRODUCTION 

The Thirty-second Expedition of the Liverpool School of 
Tropical Medicine was dispatched to Sierra Leone on November 
18th, 1914, and returned to England on April 2nd, 1915. 

The primary object of this expedition was to find a site suitable 
for the permanent research laboratory which the School proposes to 
erect in Sierra Leone. Thanks to the kindness of the Administration, 
we were enabled to approach the subject under the most favourable 
circumstances and to obtain information of the local conditions as 
regards population and incidence of disease. After other factors, 
such as healthiness, cost of erection and maintenance of the 
laboratory, and accessibility from England, had been considered, 
we were in a position to make definite recommendations which were 
embodied in a report to the Committee of the School. 

During our sojourn in the country we made a survey of the chief 
parasitic diseases of man and domestic stock, with a view to 
ascertaining the lines along which research could with most 
advantage be directed in the laboratory which it is hoped to 
establish. 

The scientific research undertaken by us deals chiefly with the 
bionomics of Glossina palpalis. The results of this and other work 
accomplished will be published in a series of papers in these Annals. 

We desire to express our deep indebtedness to His Excellency 



35 ° 


the Governor, Sir Edward Merewether, to the Principal Medical 
Officer, Dr. T. E. Rice, and to the Senior Sanitary Officer, 
Dr. R. H. Kennan, for the invariable kindness and courtesy shown 
to us during our visit to the Colony, and for the assistance rendered 
on every possible occasion. 


Some four or five miles from Freetown, at the mouth of the 
Sierra Leone River, lies the Cape Lighthouse Peninsula. This 
triangular piece of land, of which the long axis is approximately 
north and south, is separated from the mainland by Aberdeen Creek. 
On the other two sides the Peninsula is bounded by the sea. It is 
roughly three miles long, by one and a half miles broad at its base, 
with an area of about threequarters of a square mile. It tapers 
rapidly and runs as a narrow strip about ioo yards wide for nearly 
two miles, until it joins the mainland at its southern extremity. The 
Creek is fringed by a dense growth of mangroves, which in certain 
places is as much as half a mile broad. On the northern side the 
shore is rocky, and on the western side there is a gently sloping 
beach of fine white sand (see map). 

While oil palms (Eloeis guineensis ) are found all over the 
Peninsula, in certain parts they occur exclusively, the appearance of 
dense undergrowth being due entirely to the presence of innumerable 
young palms (PI. XXVII). In such places which are found especially 
in the region adjacent to the mangroves fringing Aberdeen Creek, and 
in that portion of the Peninsula on which the lighthouse and isolation 
camp are situated, there is deep shade, and little or no long grass. 
Between Aberdeen and the isolation camp the growth of palms is 
not so dense, and consequently the shade is less; in this locality 
• other trees are found, and there is a considerable amount of long 
grass. Between the isolation camp and Man of War Bay is a small 
area in which oil palms, although present in considerable numbers, 
do not predominate to the same extent as elsewhere, many other 
trees being present together with a dense undergrowth of shrub. 
Besides oil palms, cocoanut palms, mangoes and cotton trees are 
represented; there are also a few baobabs, and in the village of 
Aberdeen bread-fruit trees, guavas, plantains and golden plum 
trees. 


Annals Trop. Med. & ParasitolVol. IX 


PLATE XXVII 



Dense growth of young oil palms. 


To face p. 350. 


C. Tinlinz < 5 ^ CoLtd., Imp. 















Annals Trop. Med. £•? Parasitol., Vol. IX 


PLATE XXV111 



Hollow baobab constituting a breeding ground of Glossina palpalis. 


To face p. 351 


C. Tinling & Co., Ltd., l*f- 




35 1 


The population of the Peninsula is fairly large; in the town of 
Aberdeen are some 500 inhabitants, whilst at the Lighthouse and 
isolation camp are 50 more, and in addition there are a few natives 
living in isolated dwellings. Where the Peninsula joins the main¬ 
land is the native town of Lumley, with over 400 inhabitants. 

Goats, sheep, dogs, domestic fowls and a solitary cow were 
found, apparently in good health. Regarding the wild fauna of 
the Peninsula nothing very definite is known; monkeys 
(Cercopithecus sp.), squirrels, bats, rats and mice are numerous, as 
are also many varieties of birds, including bush fowl, guinea-fowl, 
ducks and sea birds. A few antelope, bush buck and duiker, were 
seen, but they are scarce. 

The only fresh water on the Peninsula, apart from the wells 
themselves, is the practically insignificant overflow from two or three 
springs. 

At the beginning of the dry season—December and January— 
the tsetse-fly Glossina palpalis was found in fair numbers all over 
the Peninsula. A similar prevalence of this fly is recorded by 
Dr. W. A. Young, who had spent a month in the locality during the 
previous rainy season. 

Owing to the circumscribed nature of the Peninsula, its relatively 
small size, its proximity to Freetown, and to the fact that cases of 
sleeping sickness have been found in the vicinity, the place seemed 
well adapted for studying the bionomics of Glossina palpalis. We 
were compelled to confine ourselves to one or two of the more 
important aspects of the question, as the time at our disposal was 
limited. 

Although Glossina palpalis is known to be widely distributed 
over the West Coast of Africa, and in many places the distribution 
of the fly has been carefully mapped out, but little is known of its 
pupal habitats in this region. We decided, therefore, to commence 
our investigations by a thorough search for pupae. Curiously 
enough our earliest effort in this direction was successful, as in the 
first place examined no less than eighteen empty pupal cases were 
discovered. The site (PI. XXVIII) was a large cavity in the trunk 
of a baobab tree. The cavity was conical in shape, having the ground 
as its base; the apex of the cone was some 12 to 14 feet above the 
ground. The entrance to this chamber was a large triangular 



352 


opening, about 4 feet wide at the base, which was on a level with 
the ground, and about 7 feet high at its apex; it faced south-west. 
Owing to the proximity of oil palms and a large mango, no direct 
sunlight entered the cavity through this aperture after about 
8.30 a.m. One or two small slit-like apertures, about 3 or 4 feet 
from the ground, admitted a little light to the chamber from its 
northern aspect. The floor of the cavity was, as already mentioned, 
flush with the ground, and was composed of a fine dry laterite 
gravel covered with dead leaves and debris from the inner surface 
of the tree trunk, which was ragged and rugose. The pupal cases 
were found after removal of the dead leaves, lying either superficially 
or within half an inch of the surface of the ground. Pupal cases 
were not found in the numerous crevices of the tree trunk itself, many 
of which contained earth probably carried there by ants. 

This breeding place was about six yards from the main path 
leading to the lighthouse, and about 200 yards from the latter. It 
was at least 100 yards from the sea, and not less than a quarter of 
a mile from the nearest fresh water, which consisted of a small well. 
It is interesting, and probably important, to note that pupal cases 
were only found within a foot of the trunk of the tree, and not in 
the centre of the floor of the cavity; the explanation of this peculiar 
distribution will be discussed later. 

Many other places were then searched, but with little success. 
These included the intervals between the buttresses at the roots of 
cotton trees, dark crevices in rocks, and rock surface densely shaded 
by trees, the ground immediately under and overshadowed by fallen 
trees and boulders, the cavities at the top of stumps of trees which 
had been felled, the sand along the seashore and many other places 
of a similar nature. A solitary empty pupal case was discovered on 
the sand (PI. XXIX, fig. 1) at the base of a palm tree growing on the 
margin of the sandy shore, and another was discovered lying on the 
ground in deep shade at the base of a similar tree situated in dense 
bush. 

We then decided to make a systematic search for pupae on and 
around oil palms. With this object we chose at random a young 
f palm (PI. XXXI); the tree selected was within 15 yards of the road 
and about 200 yards of the lighthouse. It was surrounded and 
overshadowed by other oil palms, which in this region formed about 




Fig. 2. General view of palm country. 


To face p. 352. 


C. Tinling <5r* Co., Ltd., Imp 


PLATE XXIX 


A finals Trop . Med. CS? Parasitol To/. / A' 



















353 


95 per cent, of all the trees present. The trunk of an oil palm 
which has its lower petioles still unremoved, as is the case with most 
of the palms in the Peninsula, is by no means easy of approach. An 
impenetrable barrier is formed by the lower petioles which project 
out horizontally in all directions to a distance of 3 feet or more from 
the trunk. Frequently the tips of the lowest petioles dip into the 
ground, an arrangement which in conjunction with the fact that the 
petioles are armed with strong sharp spines effectually prevents any 
close examination of the ground in the immediate vicinity of the 
trunk of the tree (PI. XXX and XXXI). Having first 
searched the ground beyond the range of the petioles and 
as far as possible under them without success, the petioles 
one by one were cut off at their place of origin from the trunk. 
The ground under each was then carefully searched. After 
removal of the dead leaves and light debris twenty empty pupal 
cases were discovered. The cases were found all round the 
trunk, lying, as a rule, not more than 12 inches from it; in every 
instance they were on the surface. The ground was dry and sandy, 
and covered with a certain amount of fine laterite gravel. The 
shade was dense, being derived not only from the petioles of the 
palm itself, but also from the close proximity of a large number of 
other palms, which almost completely cut off direct sunlight during 
the entire day. 

The examination of the soil around the trunks of the trees was 
greatly facilitated by the use of a spade and newspaper. A 
shovelful of superficial soil was placed on the newspaper and spread 
out in a thin layer, when the puparia could readily be seen. By this 
means the soil at different depths was more accurately and 
expeditiously examined. The superficial layer to the depth of 
about half an inch was first removed and examined, and subsequently 
the deeper layers. It was thus shown that by far the majority 
of puparia were either actually on the surface of the ground 
or less than half an inch below the surface; the deeper layers 
contained very few pupal cases. The following observation 
illustrates this point. A palm having been stripped of all its lower 
petioles, the ground around it was cleared of dead leaves and twigs. 
The soil to an extent of about one yard from the trunk was then 
removed in layers and carefully examined for puparia. The 



354 


following was the result: —The most superficial half-inch contained 
fifty-one puparia. The soil between one-half inch and three inches 
in depth contained ten, that between three inches and six inches only 
two; whilst below this depth no puparia were discovered. 

Adopting this procedure, we had not much difficulty in finding 
large numbers of puparia of Glossina palpalis in similar sites. 
Frequently between ten and twenty pupal cases were taken under a 
single palm, and on one or two occasions between twenty and thirty, 
whilst in two instances seventy-three and seventy-five puparia 
respectively were collected. 

As a result of many observations of this kind, we are enabled 
to state, as regards this Peninsula, that under any oil palm which has 
not had its lower petioles removed, which stands in dense shade and 
the ground under which is dry and not too stony, a search for the 
puparia of Glossina palpalis would in all probability be successful. 
We do not mean to imply by this that, other things being equal, 
pupae do not exist where the ground is stony, but simply that they 
are more difficult to recognise on such a surface. We put this 
postulate to the test by examining a single spadeful of superficial 
earth removed from under the petioles close to the trunk of each of 
twenty oil palms in a region where practically every tree was a palm. 
In two instances only did we fail to find the puparia of Glossina 
palpalis by this means. 

The angles which the petioles form with the trunk of the palm 
contain considerable quantities of earth and debris. These forks 
appear to be suitable for the deposition of larvae, and accordingly 
a very careful search for puparia was made. Before examining the 
contents of the angles, all the pupae lying on the ground to a depth 
of three inches were first removed with the aid of a spade, care being 
taken to disturb the petioles as little as possible. The ground 
around the trunk was then covered with newspapers and the petioles 
stripped from the tree, and all the soil and debris in the angles 
collected and examined. In this manner several palms were entirely 
stripped of petioles, whilst odd petioles were removed from many 
other palms (PI. XXXI and XXXII). The result was striking, 
a solitary empty pupal case only being discovered in this situation. 

At the base of other trees, especially cotton trees and mangoes, 
are what appear ideal pupal habitats from the point of view of shade 
and soil. The result of searching these places was, however, 




Annals Trop. Med. & ParasitolVol. IX 


PLATE XXXI 



Oil palms ; the lower petioles have been stripped from the tree on the left. 


To /ace p. 354 . 


C. T in ling Co., Ltd., Imp . 





Annals Prop. Med. & Parasitol., Vol. IX 


PLATE XXXII 



Young oil palm from which all the petioles on the right side had been stripped. 


To face p . 354 . 


C. Tittling <5r* Co., Ltd., Imp. 





PAGE NOT 
AVAILABLE 













355 

decidedly disappointing, only an occasional puparium being 
discovered. 

As might be expected, most of the puparia taken were empty. 
Out of a total of about 450 only twenty were unhatched.* From a 
number of these Glossina palpalis emerged after they had been kept 
in the laboratory some days. 

We have already referred to the fact that the Aberdeen Creek 
side of the Peninsula is fringed with extensive mangrove swamp. It 
is popularly supposed that there is some connection between the 
presence of these swamps and the occurrence of Glossina palpalis. 
In fact, the local name for the ‘ fly ’ amongst the natives is ' mangrove 
fly.* That portion of the swamp in the vicinity of Aberdeen village 
was carefully examined by us firstly for Glossina palpalis , and 
secondly for its pupae. It should be mentioned that while the roots 
of the mangroves are covered at high tide to a depth varying from 
a few inches to three or four feet, at low tide the sea recedes to a 
considerable distance beyond their outer fringe, leaving amongst 
them small collections of water and tiny streams, so that the 
mangroves are seen to be growing in soft sandy mud and pools of 
water (PI. XXXIII). An inspection of the mangroves was made on 
severed occasions, both at low and high water. The outer edge was 
examined on foot at low water, but a boat was necessary at 
high tide. 

Glossina was found not only in the midst of the mangroves, but 
also at their outermost fringe, both at low and high water; this is 
of considerable interest, as at certain points where the tsetse was 
seen the distance from the dry land was at least half a mile. No 
trees were found growing amongst the mangroves below high water, 
although just above this point a few mangroves were noticed inter¬ 
mingled with the ordinary bush growth. The mangroves themselves 
afforded fairly good shade, and this, together with the fact that a 
considerable number of paths were cut through them to the fishing 
grounds beyond, and that the mangroves are inhabited by large 
numbers of small birds, mud fish, crabs, shellfish, snails, wading 
birds, etc., which may afford a supply of food for the tsetse, is 
probably sufficient reason for its presence. 

The next question to be considered is, do the mangroves 

* The extreme length and breadth of 16 unhatched pupae were measured. The following is 
a summary of the results :—Length : max. 6*i mm., min. 5*6 mm; average 5*93 mm. Breadth : 
max. 3*6 mm; min. 3*0 mm ; average 3*25 mm. 



356 


constitute a breeding ground for Glossina palpalis ? Bagshawe 
(1911) writes: ‘It is probably lost labour to look in moist soil and 
mud for pupae of tsetse flies; they have never been found in any 
situation which is not at least moderately dry.* Nevertheless, in 
view of the fact that the fly was found amongst the mangroves, we 
determined to investigate the point carefully. At low tide, with the 
aid of six assistants, all of whom had had considerable experience 
in finding pupae, we made an exhaustive examination of the sandy 
mud amongst the mangroves and also of the mangroves themselves. 
The latter, however, afford very few possible places for the lodgment 
of pupae, as the stems are smooth and narrow and do not present 
many forks or clefts in which pupae could lodge. Similar examina¬ 
tions were made at high tide over the same area. No puparia were 
found on the mud or floating on the water amongst the mangroves, 
nor were any discovered on the stems and branches. 

As a result of this work, it appears clear that the tsetse which we 
observed in the mangrove swamps were not breeding there, but had 
travelled from the land. Support to this conclusion was furnished 
by examination of the bush immediately above high-water mark. 
Puparia were found there under oil palms in large numbers, and a 
few under other trees. Apart from the failure to discover puparia in 
the mangrove swamps such places do not appear to be suitable 
breeding grounds for tsetse-fly, as pupae which are deposited on 
sea-water or on ground which is covered by sea-water for a certain 
number of hours each day do not hatch, as is shown by the following 
experiment: — 

Experiment. —Eighteen unhatched pupae found at the base of oil palms 
were placed in glass jars containing a small quantity of sand. Nine of the jars 
were half filled with sea-water for a period of three hours daily during a month. 
The pupae were observed to float horizontally on the surface of the water, the 
tubercles and stigmata being as a rule just below the surface. When the water was 
decanted at the end of the three hours, the pupae were left lying on the moist 
sand. This experiment was an attempt to imitate the conditions to which pupae 
deposited in mangrove swamp would be subjected, except that in the experiment 
they were exposed to water once a day only, whereas in nature the tides cover the 
mangrove swamps twice daily. At the end of a month the daily addition of water 
was discontinued and the pupae were left lying in the sand which soon became 
dry. No water was added to the other nine jars, the pupae in which served as 
controls. Six of the nine control pupae hatched within the time during which 
the experiment was continued, viz. : 35-60 days, depending on the date when the 
pupae were discovered. The flies emerged on the 1st, 10th, 20th, 23rd, 27th and 
32nd days respectively after the commencement of the experiment. No flies 
emerged from the pupae which had been exposed to sea-water. The temperature 
was that of the laboratory 8o°-86° F. 




357 


Carpenter's (1912) work on this subject is of great interest; he 
writes ‘According to my experience pupae are only found in the 
driest possible situation, though always close to the water’s edge; 
so that it would appear that contact with water is inimical to them. 
It will be seen, however, that they have a considerable capacity to 
resist the effects of submersion; moreover, they do not become mouldy 
(so long as they are alive) when kept in an atmosphere with the 
maximum degree of humidity. Their powers of resistance to sun 
are less in proportion.* He concludes from experiments that should 
any breeding ground be flooded at intervals by heavy rains the 
conditions would not completely destroy all the pupae. Even four 
successive submersions for twenty-four hours destroyed only half the 
number. Further experiments showed that it is rather the frequency 
of submersion than the total time that is adverse to the pupae. He 
states that a certain proportion can survive flotation for eight days, 
but none for ten days. 

A general survey of the pupal habitats discovered by us in the 
Cape Lighthouse Peninsula reveals several important facts. 

Firstly, fly and pupae are distributed more or less evenly over 
the whole extent of the Peninsula. They are certainly not limited 
to the immediate vicinity of water, both being found at least a 
quarter of a mile from the seashore. This is the more remarkable 
in view of the almost complete absence of fresh water. There is, 
so far as we are aware, no record in the literature of palpdlis pupae 
being found in any situation except on the water's edge. Bagshawe 
(1908), who was the first to discover the pupae of Glosstna palpalis 
in nature, writes ‘ One may for the present say that the larvae are 
dropped in shade, it may be of shrubs, it may be of bananas, within 
forty-five yards of water.' Fraser and Marshall (1909) state that 
the distance from high-water mark at which deposits were found was 
never more than fifteen yards, the usual distance being five yards; 
whilst Carpenter writes * According to my experience pupae are only 
found in the driest possible situations though always close to the 
water's edge.’ It must be noted, however, that these statements 
refer to East Africa. The observers quoted worked on Lake 
Victoria Nyanza, where conditions appear to be rather different from 
those obtaining in Sierra Leone. Whilst enormous numbers of flies 
are to be found in the former place, and they appear to be strictly 



358 


limited in their distribution to the lake shore and larger rivers, as 
regards the Cape Lighthouse Peninsula this is certainly not the case. 
Glossina palpalis is not nearly so numerous, nor is it limited to the 
immediate vicinity of water. 

Secondly, pupae were concentrated on the earth close to the 
trunks of trees, and were not found to any extent on the trees 
themselves or on the ground between them. By far the majority of 
the puparia taken by us were less than one foot from a tree trunk. 
The explanation of this appears to be that the tsetse deposits its 
larvae in the most secluded and shady spot available. Possibly the 
larva is deposited whilst the fly is resting on the trunk or on the 
under surface of a lower petiole. Such a hypothesis would account 
for the peculiar distribution of the puparia in the hollow baobab 
referred to previously. Tsetses resting on the inner surface of the 
hollow trunk would drop their larvae on the ground close to the tree 
trunk, thus explaining the peripheral distribution of the puparia on 
the floor of the chamber. Similarly in the case of the oil palm, flies 
resting on the under surface of the lower petioles or on the tree trunk 
would also drop their larvae on the ground near the trunk. 

The facts discovered by Moiser (1912) are of interest in this 
connection. He put men in trees at a height varying from 10 to 
25 feet for an hour; not one of them reported having seen a fly, 
though several ( Glossina tachinoides') were seen on the ground during 
the period. In order to determine where flies usually rest, he put 
up a large mosquito net (8 x 8 x 8 ft.) in the bush, thus enclosing 
a portion of the natural haunt of the fly. Eleven Glossina 
tachinoides were then liberated within the net. A number of them 
quickly disappeared from view, and after a few minutes* search he 
discovered them resting in an inverted position on the under side of 
small branches and twigs close to the ground. Branches between 
6 inches and 2 feet from the ground were those most frequently 
occupied by the tsetses, and in order to observe them closely Moiser 
had to lie on the ground. Subsequently flies were found at rest on 
the under surface of small branches outside the net. 

Thirdly, the ground at the base of oil palms appears to be a most 
suitable breeding place for Glossina palpalis. In this region 
puparia of Glossina palpalis can be found under almost any oil palm 
which has not had its lower petioles removed, which stands in dense 



359 


shade, and the ground under which is dry and not too stony. 
Moreover, Glossina palpalis is able to breed in a locality where oil 
palms of various sizes and ages 'afford the sole shelter. The lower 
surface of a petiole is broad and smooth and well protected from 
the sun, and tsetse have been observed in this situation. A fly in 
this position is hidden from view except to anything actually on the 
ground, and is also to an extent protected from molestation by the 
sharp strong spines with which the petioles are armed. In so far 
as the Cape Lighthouse Peninsula is concerned, the situations in 
which pupae are deposited are hidden from view, probably as a 
protective measure for the fly undergoing parturition and for the 
newly deposited larva against birds, a view which would explain 
the absence of puparia in many spots, such as the spaces between 
the buttresses of the trunks of cotton trees, otherwise apparently 
well adapted as pupal habitats. 

Zupitza (1908), working at Duala in the Cameroons, recorded 
that he had found the puparia of Glossina palpalis in the humus and 
moss in the forks of branches and in the cracks in the bark of all 
trees, especially in the angles of leaf sheaths of palms at a height of 
a few centimetres to three and a half metres above the ground. He 
did not look for them at a higher level. They were never found in 
the dry mould of hollow trees, nor in or on the ground. The 
observations of Zupitza are in curious contradistinction to those 
made by us. The first pupal habitat discovered by us was the 
ground forming the floor of the chamber in the baobab trunk. 
Moreover, whilst we constantly found puparia on the ground close 
to the trunk of oil palms, only a single empty case was discovered 
in the angles between the petioles and the tree trunk. As already 
mentioned, we most carefully investigated this place as a possible 
pupal habitat, entirely stripping the petioles from three or four 
palms on the ground under which large numbers of pupae had 
been found, and examining the debris between them and the tree 
trunks without finding any pupae. In addition, odd petioles were 
stripped from many other palms with equally negative results. 

The explanation of this apparent discrepancy is cryptic. We are 
not able to suggest any reason why Glossina palpalis should in one 
locality deposit its pupae on the ground and in another on the tree 
itself, unless in the latter case the ground happened, through 



3 < 5 o 


moisture or some other unknown condition, to be an unsuitable pupal 
habitat. Zupitza states that in the angles between the petioles and 
the trunks of the palms the pupae would be kept moist, but that they 
would be in no danger of being drowned or washed away by rain¬ 
water running down the trunk. He is evidently under the 
impression that some amount of moisture is necessary, as he never 
found them in the dry moss in the holes of trees, nor in or on the 
ground. This is not our experience, as they were almost invariably 
found lying on or just under the surface of perfectly dry ground. 

Before leaving this subject, it is of interest to consider what steps 
it would be necessary to take in order to clear the Peninsula of 
tsetse. In view of the fact that the fly is ubiquitous in the area under 
consideration and that its pupae have a like distribution, it is quite 
obvious that no localised or partial clearing could be recommended 
as likely to eradicate the fly. Moreover, in districts where practi¬ 
cally all the vegetation consists of oil palms, wholesale removal of 
these cannot, for economic reasons, be contemplated. Yet we are 
faced with the fact that it is under the young palms that the tsetse- 
fly find their best breeding place; any method to be practicable 
must not interfere with the economic value of the trees. The point 
that has to be considered, therefore, is whether any measures 
short of actual destruction of the trees would suffice to 
get rid of the fly. In our experience the ground at the 
base of an oil palm which has had its trunk cleared of petioles 
does not constitute a breeding place of palpalis. The soil 
at the base of many such trees was searched without success. 
We are convinced that it is the presence of the lower petioles that 
renders young oil palms such excellent breeding places. Removal 
of these would, in our opinion, destroy the breeding grounds of 
palpalis , and by this means such localities could be freed of fly 
without loss by damage to the trees. Such a procedure would be 
laborious, but it must be remembered that the petioles when once 
removed do not grow again. In old palms the lower petioles are 
absent; as the tree grows the lower, that is the older, petioles become 
dry and rotten and are easily stripped off, but in very young 
palms they are much tougher, and hence more difficult to remove. 
Naturally, in those places where shrub growth, other than young oil 
palms, occurred, it would have to be removed. 



361 

It may not be considered advisable on economic or epidemio¬ 
logical grounds to attempt to rid the Peninsula of tsetse at the 
present time. If, however, this can be undertaken as an experiment, 
results of the utmost value would be obtained, and the information 
gained could be applied in districts where for economic or 
epidemiological reasons it might be of vital importance to exterminate 
Glossina palpalis. Owing to its proximity to Freetown, its small 
size and its well circumscribed character, the Cape Lighthouse 
Peninsula is eminently suitable for an experiment of this nature. 


CONCLUSIONS 

1. The breeding grounds of Glossina palpalis are not so 
strictly limited to the immediate vicinity of water as has hitherto 
been thought; they may occur quite independently of fresh water 
and at least a quarter of a mile from sea water. 

2. Although Glossina palpalis is to be found in considerable 
numbers in mangrove swamps and may travel in these to a distance 
of at least half a mile from dry land, the swamps do not constitute 
a breeding ground of the fly. 

3. The pupae of Glossina palpalis do not hatch when subjected 
to daily flotation on sea water. 

4. The ground around the trunk of oil palms ( Eloeis guineensis) 
which have not been stripped of their lower petioles constitutes an 
excellent breeding place for Glossina palpalis . 

5. Glossina palpalis can breed in localities in which practically 
the only tree is the oil palm. 

6. Stripping the oil palm of the lower petioles would suffice to 
destroy the breeding ground in such localities. 



REFERENCES 


Bagshawe, A. G. ( i 911). Sleeping Sickness Bulletin , Vol. Ill, p. 362. 

- (1908). Reports of the Sleeping Sickness Commission of the Royal Society , No. IV, p. 48. 

Carpenter, G. D. H. (1912). Progress Report of Investigations into the Bionomics of Glossina 
palpalis , July 27, 1910, to August 5, 1911. Report of the Sleeping Sickness Commission of 
the Royal Society } No. XII, pp. 79-1 n. 

Frazer and Marshall (1909). Progress Reports on Uganda Sleeping Sickness Camps. London 
Sleeping Sickness Bureau , pp. 40-44. 

Moiser, B. (1912). Notes on the haunts and habits of Glossina palpalis. Bull. Ent. Res Vol. 
Ill, pp. 195-202. 

Zupitza, — (1908). Ueber die Schlafkrankheitsfliege bei Duala. Beibefte zum Arcbiv. f. Scbiff. 
und Tropen. Hyg. y B. 12, pp. 1-27. 





J63 


FOOD OF GLOSSINA PALPALIS IN 
THE CAPE LIGHTHOUSE PENINSULA, 

>> SIERRA LEONE 

[Being the Second Report of the Thirty-second Expedition of the 
Liverpool School of Tropical Medicine, 1914-1915.] 

BY 

WARRINGTON YORKE 

AND 

B. BLACKLOCK 

(Received for publication 28 April , 1915) 

Although there is a certain amount of evidence to show that 
Glossina palpalis can take up water and vegetable juices, practically 
all observations indicate that for its continued existence vertebrate 
blood is necessary. In view of the small amount of time at our 
disposal, and of the comparative scarcity of Glossina palpalis , we 
were able to examine only 200 flies in respect of the nature of the 
blood found in their intestinal tract. The tsetse were dissected 
immediately they were brought into the laboratory. As at the same 
time the flies were examined for the presence of trypanosomes, they 
were dissected in the manner described by Lloyd (1912). Briefly 
it consists in splitting the dorsum of the thorax longitudinally with 
a cataract knife, and then drawing out the salivary glands by gentle 
traction on the head. Usually the glands are removed intact 
attached to the head, the oesophagus breaking off at the point where 
it enters the pharynx; occasionally they break across when only 
partially withdrawn, but in this case are easily caught up with a 
fine pair of forceps. The proventriculus is then sought at the ventral 
surface of the thorax, and is drawn out with the sucking stomach 
and most of the intestine. The posterior portion of the hind-gut 
and rectum are removed by cutting off the last segment of the 
abdomen. The gut contents and those of the salivary glands and 
of the proboscis were examined between a slide and coverslip in a 
fresh unstained condition. 



364 


Recognisable red blood cells were seen in 16 of the 200 flies thus 
examined. It is necessary to point out here that only those flies in 
which definite red blood corpuscles were seen are recorded as 
containing blood. Practically every fly had in its mid-gut a certain 
quantity of pigmented material, the colour of which varied from 
bright red to brown or black. Such pigmented material does not, 
however, necessarily imply the presence of red blood corpuscles; on 
microscopical examination it was usually found to consist of granular 
debris and globules appearing red or brown in colour, and 
crystals which in many cases were probably haemin or haemoglobin 
crystals. Although it is highly probable that this pigmented 
material was derived from blood, still one could not be certain, and, 
therefore, in the following table only those flies in which definite 
red blood corpuscles were encountered are considered as containing 


blood. 

Tablx I.—Result of examination of freshly-caught wild Glonima fslptlts for recognisable blood 

corpuscles 



Number 

examined 

Number in which 
recognisable 
mammalian red 
corpuscles were seen 

Number in which 
recognisable 
nucleated red 
corpuscles were seen 

Males . 

113 

8 

0 

Females . 

! 

87 

6 

2 

I 

Total . 

200 

14 

2 


From the limited number of observations made, it appears that 
eight per cent, of freshly caught Glossina palpalis in this Peninsula 
contain recognisable red blood corpuscles—mammalian blood in 
seven per cent., and nucleated red cells of an undetermined nature 
in one per cent. The chief source of mammalian blood on the 
Peninsula is probably man and his domestic stock—goats, sheep, 
and dogs. In addition there are a few antelope—bushbuck and 
duiker were seen—and a considerable number of monkeys 
(<Cercopithecus sp.) squirrels and bats, rats and mice. It is 
interesting to note that the mammalian red cells seen in the fourteen 
flies observed to contain these, were of the large type and readily 
distinguishable from the small variety found in sheep and goats. 





36s 


The source of nucleated blood corpuscles is apparently much 
larger. Birds and lizards are numerous, and in the mangrove 
swamps there are vast numbers of mud-fish, crabs, shell-fish, snails, 
wading birds and sea birds. No crocodiles occur. In view of the 
fact that the reservoir of nucleated blood corpuscles is probably much 
greater than that of non-nucleated red cells, it is interesting to find 
mammalian blood in fourteen Glossina palpalis and nucleated red 
cells in only two, an observation which appears to indicate that 
Glossina palpalis either prefers mammalian blood or finds it more 
easy to obtain. The flies were captured at various parts of the 
Peninsula, but owing to its small size none of these localities was 
more than a quarter of a mile from human habitation—either the 
village of Aberdeen or the settlements at the isolation camp and the 
Cape Lighthouse. 

In order to appreciate the real meaning of the results obtained 
by such an examination of the gut of freshly caught tsetse in respect 
of blood corpuscles, it is of importance to have at our disposal some 
data regarding the length of time red blood cells can be recognised 
in the intestine of a fly after feeding. To obtain this information 
freshly caught tsetse were fed on a rat or fowl and dissected after 
various intervals, and the gut contents examined for the presence of 
red blood cells. The results are given in the Tables II and III. 

In the case of mammalian blood it is seen that in over 90 per cent, 
of the flies red cells can be recognised 24 hours after a feed, but in 
only 40 per cent, of those examined after 48 hours, whilst in no 
instance were definite red cells noted after 72 hours. The fowl red 
blood cells were found to be recognisable for longer periods; after 
24 hours 100 per cent, of the flies showed nucleated red blood cells, 
whilst after 48 hours red blood cells were seen in 60 per cent., and 
after 72 hours in 40 per cent. The temperature was that of the 
laboratory, 8o°-86° F. Possibly to a certain extent this difference 
is to be explained by the fact that the nucleated red blood 
cells of the fowl are more characteristic, and hence more easily 
recognised than are the non-nucleated red cells of mammals. Of 
course, these figures are approximate, as only 64 tsetse were 
employed in the experiment: they are, however, sufficiently accurate 
for practical purposes, although some observers have recorded, in the 
case of isolated flies, that blood could be found after much longer 



3 66 


intervals. Thus in the case of Glossina morsitans , Lloyd (1913) 
observed fowl red cells in a clot in the sucking stomach several 
weeks after the fly had last fed on a fowl, monkeys having been used 
as blood donors in the interval. Nevertheless, these figures show 


Table II.—Experiment to ascertain the length of time rat red blood corpuscles can be recognised in 

G. palpalis 


No. 

Date fed 

Date 

examined 

No. of 
hours 
after 
feed 

Sex 

Result of 
examination. 
Recognisable 
blood cells 
present or absent 

Percentage of flies 
in which 
blood was found 

1 


18.12.14 

24 

<5 

+ + + 



2 

17.12.14 

18.12.14 

2 4 

(J 

+ + + 



3 

17.12.14 

18.12.14 

24 

$ 

+ + + 



4 

17.12.14 

18.12.14 

24 

9 

+ + + 



5 

17.12.14 

18.12.14 

24 

9 

+ + + 



6 

17.12.14 

18.12.14 

24 

3 

+ + 4 - 


9 2 

7 

17.12.14 

18.12.14 

24 

9 

4 - + + 



8 


18.12.14 

24 

<? 

4 - 4 - 



9 


2 9- I * , 5 

24 

<? 

— 



10 


2 9* I * I 5 

24 

<? 

4 - 4 - 4 - 



11 

9 

3* 2 * I 5 

24 

<? 

4 - 4 - 4 - 



12 

2.2.15 

3- 2 * , 5 

24 

<? 

4 - 4 - 4 - 



*3 

28.1.15 


48 


_ 




28.1.15 


48 

9 

— 



*5 

28.1.15 


48 

<? 

4 - 4 - 4 - 



16 

28.1.15 

1 

48 

9 

— 



l 7 

28.1.15 


48 

9 

+ 4 - 4 - 


40 

18 

2.2.15 


48 

9 

— 



*9 

2.2.15 


48 

<j 

— 



20 

2.2.15 


48 

9 

+ 



21 

2.2.15 

. 9 

48 


— 



22 

2.2.15 

4* 2 * , 5 

48 

<j 

+ + + 



2 3 

28.1.15 

3 »»>S 

7 2 


— 



2 4 

28.1.15 

3 I-I-I 5 

7 2 

cf 

— 



2 5 

28.1.15 

31.1.15 

7 2 

9 

— 



26 

2.2.15 

5 * 2 * 5 

7 2 

9 

— 



2 7 

2.2.15 

5- 2 *5 

7 2 


- 



28 

2.2.15 

5- 2I 5 

7 2 

<? 

— 


0 

2 9 

2.2.15 

5 * 2 - 5 

7 2 

9 

— 



30 

2.2.15 

5- 2I 5 

7 2 

9 

— 



3 * 

2.2.15 

5- 2, 5 

7 2 

<? 

— 



3 2 

2.2.15 

5* 2<I 5 

7 2 

9 

— 



33 

2.2.15 

5- 2I 5 

7 2 


— 



34 

21.1.15 

2 5 .i.i 5 

96 

<? 





that as a general rule mammalian (rat) red blood cells are no longer 
recognisable after a period of 72 hours, whilst fowl red cells are 
observed in only 40 per cent, of cases after a similar period. This 
information is of importance in connection with that obtained by 















3 67 


examining the blood found in freshly caught tsetse-fly. When the 
statement is made that only seven per cent, of such flies contain 
mammalian blood, it must be borne in mind that if the flies had all 
fed on a mammal 48 hours previously, we would expect to find 
red cells in only 40 per cent., whilst if they had fed 72 hours 
previously very few would contain recognisable red cells. 


Tablk III.—Experiment to ascertain the length of time fowl red blood corpuscles can be recognised in 

G. palpalis 


No. 

Date fed 

Date 

examined 

No. of 
hours 
after 
feed 

Sex 

Result of 
examination. 
Recognisable 
blood cells 
present or absent 

Percentage of flies 
in which 
blood was found 

1 

28.1.15 

29.1.15 

24 

3 

+ + + 

'I 

2 


3 II - , 5 

24 

3 

+ + + 


3 

30.1.15 

3 ,II 5 

24 

6 

+ + + 


4 

30.1.15 

3 III 5 

24 

3 

+ + + 


5 

I.2.I5 

2.2.15 

24 

9 

+ 

100 

6 

I.2.I5 

2.2.15 

24 

3 

+ + + 


7 

I.2.I5 

2.2.15 

24 

? 

+ + + 


8 

I.2.15 

2.2.15 

24 

3 

+ + + 


9 

1.2.1 5 

2.2.15 

24 

3 

+ 


10 

I.2.I5 

2.2.15 

24 

9 

+ + 


11 

28.I.I5 

30.1.15 

48 

3 

_ 


12 

28.1.15 

30.115 

48 

? 

— 


! 3 

28.1.15 

30.1.15 

48 

3 

— 


H 

28.I.I5 

3°* I * I 5 

48 

3 

+ + + 


i 5 

28.I.I5 

30.1.15 

48 

$ 

+ + + 

60 

16 

1-2.15 

3.2. 1 5 

48 

3 

+ 


l 7 

I.2.I5 

3.2.15 

48 

3 

+ + + 


18 

I.2.I5 

3 - 2-15 

48 

3 

— 


>9 

I.2.I5 

3 - 2 -i 5 

48 

3 

+ + + 


20 

I.2.I5 

3.2.15 

48 

3 

+ + + 


21 

28.I.I5 

3 i-i-i 5 

72 

3 

— 


22 

28.1.15 

3 i-i-i 5 

72 

3 

+ 


23 

28.1.15 

3 i-i-i 5 

72 

? 

+ 



28.1.15 

3 i-i-i 5 

72 

3 

- 



28.1.15 

3 i-i-i 5 

72 

9 

— 

40 

26 

30.1.15 

2.2.15 

72 

3 

— 


27 

30.1.15 

2.2.15 

72 

3 

+ + + 


28 

30.1.15 

2.2.15 

72 

9 

— 


29 

30.I.I5 

2.2.15 

72 

3 

+ + 


30 

30.1.15 

2.2.15 

72 

3 




The question whether tsetse-fly take up other food than blood 
is one which is difficult to decide. Stuhlmann (1907) and Degen 
(1909) came to the conclusion that they did not, but Maugham (1911) 
states that he has seen tsetse-flies sucking vegetable juices on two 
occasions. In 1905 he observed a Glossina morsitans alight on a 






368 


stem of young marsh grass ( Phragmites communis) and deliberately 
insert its proboscis and unmistakably suck for a period of about 
three and a half minutes. At this stage Maugham caught the fly, 
and found on examination that it was partly full of the moisture 
from the plant. Again in 1908, in an absolutely gameless and 
practically waterless country, he observed Glossina morsitans feeding 
on a piece of sugar-cane at a point where the pith was exposed. He 
attempted to catch this fly, but unfortunately failed to do so. 
Taute (1912) investigated the point experimentally in the following 
manner. Two hundred tsetse-flies which had been fed daily on 
blood for a period of two months were starved for five days, and 
subsequently were given an opportunity of feeding on small pieces of 
mango fruit. Taute records that in three cases Glossina morsitans 
buried its proboscis completely in the fruit and remained in this 
position for several minutes, but definite sucking did not take place. 
One of the flies was killed and dissected immediately after this 
operation. Neither in the lumen of the proboscis nor in the 
oesophagus or remainder of the alimentary canal could the smallest 
trace of mango juice be recognised. 

The work of Carpenter (1912-13), however, affords strong support 
to the view that Glossina palpalis does take up other food than 
blood. Carpenter examined the intestinal contents of a large number 
of freshly caught G . palpalis , and found in a proportion of 
them small fragments of tissue of an obviously vegetable origin, 
e.g., pieces of vegetable parenchyma, starch grains, pieces of alga, 
and a minute fungus. 

Apart from the solitary instance recorded by Maugham, there 
appears to be no direct evidence that tsetse-fly will imbibe anything 
other than vertebrate blood, and, moreover, the evidence offered by 
him is by no means convincing, as he omits to state the manner in 
which he examined the fly or how he detected that it was partly 
filled with moisture from the plant. Stuhlmann affirms that Glossina 
lives exclusively on living blood, and that they refuse to take up 
shed blood, water or syrups; and Degen attempted to feed them on 
fruit, saccharine fluids, meat, etc., without success. Rodhain, 
Pons, Vandenbranden and Bequaert (1912) observe that it is 
generally admitted that tsetses cannot engorge themselves with 
blood unless they obtain it directly from the capillaries in which the 


369 


fluid is maintained under a definite pressure. Leaving pressure 
out of the question for the moment—Rodhain and his collaborators 
have since shown that this is not an essential factor—all experimental 
evidence shows that Glossina are unable to imbibe shed blood. In 
the experiments described below we have demonstrated that they will 
not absorb certain other fluids offered to them in open vessels. 

Fourteen flies which had been starved for 48 hours were placed in 
each of two cages. In one of these was a petri dish filled with the 
following solution:— 

Sodium chloride. 0*9 g. 

Sugar . 2 - og. 

Neutral red . o ig. 

Water . 100 c.c 

and in the other a petri dish containing the following: — 

Methylene blue. 0*05 g. 

Sodium carbonate . 0*025 S- 

Water . 100 c.c. 

Small twigs and leaves were floated on the surface of the solutions. 
The flies were dissected as they died; all those that were still alive 
at the end of 48 hours were killed and examined. There was no 
indication that any of the flies had imbibed fluid. These results 
are conclusive because, as will be seen later, solutions of neutral red 
and of methylene blue in the concentrations used, if taken up by 
Glossina, stain the tissues densely. Such work as we have done, 
therefore, confirms the generally accepted view that Glossina is 
unable to imbibe exposed fluids directly. 

The work of Rodhain and his collaborators showed, however, 
that tsetses can suck up citrated blood through a membrane consisting 
of the freshly removed skin of a mouse. We made use of this 
observation to prove that not only will tsetses take up citrated blood 
through such a membrane, but that they will also imbibe various 
other fluids. The apparatus used by us was a slight modification of 
that figured and described by Rodhain. It consisted of a short glass 
cylinder about three-quarters of an inch in diameter, the lower end 
of which was closed by a cork through which the shorter limb of an 
b-shaped piece of glass tubing passed; the upper end of the 
cylinder was covered by the membrane. The pressure of the fluid 
in the cylinder was indicated by its level in the longer limb of the 
glass tube. 



37 ° 


In our first series o£ experiments flies were fed on defibrinated 
goat-blood through a membrane of fresh rat’s skin. It was observed 
that (within the limits of the experiment) the pressure made no 
difference to the manner in which the flies engorged themselves. 
They distended themselves without the least difficulty at pressures 
ranging between + or — 60 mm. of blood. It was further noted 
that full distension of the flies occurred at least as rapidly and 
regularly as when the flies were allowed to feed on live rats. On the 


Table IV.—Giving the results of feeding G. palp alts on defibrinated blood and various dilutions of this with 
normal salt solution, through a* membrane consisting of rat skin 


No. 

Sex 

Membrane 

Nature of fluid 

Result 

Remarks 

, 

<? 

Rat skin . 

Undiluted goat blood 

Complete distension 

Immediately 

2 



n 

it 

11 

3 

9 


a 

a 

a 

4 

9 

11 

a 

it 

11 

5 

3 

it 

a 

ii 

a 

6 

9 

M 

a 

ii 

a 

7 

<? 

II 

50% goat blood 

it 

a 

8 

3 

it 

n 

it 

a 

9 

9 

it 

n 

a 

a 

IO 

3 

» 

25% goat blood 

a 

a 

ii 

9 

n 

i) 

it 

a 

12 

3 

it ••• ••• 

a 

a 

a 

*3 

3 

11 

10 % £ oat blood 

a 

a 

*4 

9 

a 

n 

Partial distension ... 

After 15 minutes 

>5 

9 

tt 

a 

11 

i» 

16 

3 

a 

10% rat blood 

Complete distension 

Immediately 

*7 

3 

a 

a 

a 

11 

18 

3 

a 

a 

a 

11 

19 

3 

a 

5% goat blood 

No visible distension 

After 15 minutes goat*' 
cells seen in anterior gut 

20 

9 

a ••• 

11 

1* 

After 1 5 minutes no goats’ 
cells seen in anterior gut 

21 

9 

a 

5 % rat blood. 

11 

After 15 minutes rats’ cell* 
seen in anterior gut 

22 

9 

jj ••• ••• 

n 

Partial distension ... 

After 15 minutes 

^3 

3 

ii 


No visible distension 

After 15 minutes red cells 
seen in anterior gut 

24 

3 

ii 

a 

Marked distension ... 

After 15 minutes 


other hand the flies attacked the membrane with less alacrity than 
they do the skin of the living animal. Nevertheless, many flies 
settled on and pierced the membrane at once, and, as a rule, a little 
patience was sufficient to induce a large proportion of the flies to 
feed. An experiment was conducted to determine the effect of 
diluting the defibrinated blood with salt solution in various degrees; 
the results are given in Table IV. It was found that Glossina palpalis 













37i 


engorged itself with the following dilutions, viz., 50 per cent, blood, 
25 per cent, blood, and 10 per cent, blood, but the dilution 
consisting of 5 per cent., blood and 95 per cent, salt solution was 
not taken up by the tsetses with the same rapidity and regularity 
as the other solutions. Of the six flies which were offered the last 
dilution of blood, only two partially distended themselves; in the 
other four no distension was observed, but in three of these red ™»11c 
were seen on dissection. 

Physiological salt solution alone was then tried, but although the 
flies attacked the membrane with the same eagerness as before, no 
visible distension was noted, except in one instance where the fly 
appeared to become partially engorged. Tsetses frequently inserted 
their proboscis and appeared to endeavour to feed; in some 
instances the membrane was pierced probably at least 100 times with 
apparently no result. We shall return later to the question whether 
these flies had actually taken up any salt solution, and simply 
note here that no distension comparable to that observed when 
deflbrinated blood is offered took place. 

We next turned our attention to the question of what element of 
the blood proves so attractive to the fly that it engorges itself with 
this fluid to such a marked degree. Fresh deflbrinated goat-blood 
was centrifugalised, and the red cells separated from the plasma; 
the red cells were then washed free from plasma with normal sodium 
chloride solution and a 50 per cent, suspension of them made in 
salt solution—a concentration which corresponds approximately to 
that of red cells in normal goat-blood. It was found that Glossina 
palpalis engorged itself readily with the red cell suspension. By the 
addition of normal salt solution, suspensions containing 25 per cent., 
10 per cent, and 5 per cent, of red blood cells were made. The 
results of offering these to the flies are given in Table V. The tsetse 
engorged themselves completely with all the higher concentrations, 
but in the case of the 5 per cent, suspension of red blood cells only 
one of three flies became completely distended, whilst in the other 
two no distension was observed, although red cells were found in the 
gut on dissection. 

Attempts were made to feed a number of Glossina palpalis on 
fresh deflbrinated plasma. The plasma was obtained by shaking 
up goat’s blood in a bottle with beads. After separation of the fibrin 



372 


by straining through gauze, the red cells were thrown down by 
centrifugalisation and the plasma siphoned off. Plasma thus 
prepared is always of a slightly reddish tint, owing to a certain 
amount of damage to red cells during the process of defibrination, 
but the amount of haemoglobin dissolved is so small as to be 


Tabu V.—Giving the results of feeding G. palpalis on suspensions of red blood cells of various concentration 

through a membrane of rat skin 


No. 

Sex 

Membrane 

Nature of suspension 

Result 

Remarb 

i 

(? 

Rat sldn . 

5° % goat erythrocytes 

Complete distension 

Immediately 

2 

? 

» 

ii 

11 

i> 

3 

<j 

» ••• 

ii 

11 

11 

4 

<? 

n 

ii 

n 

1? 

5 

<j 

ii ... ... 

ii 

11 

11 

6 

$ 

n 

ii 

n 

11 

7 


,, . 

25 % goat erythrocytes 

11 

11 

8 

<? 

11 

1* 

ii 

11 

9 

<? 

11 

i° % goat erythrocytes 

Partial distension ... 

After 15 minutes 

IO 

$ 

11 ••• 

>» 

>» 

s ii 

ii 

c 1 

„ . 

u 

Complete distension 

Immediately 

12 

<? 

„ . 

5 % goat erythrocytes 

i> 

After 15 minutes 

>3 

9 

11 ... ... 

11 

No visible distension 

After 15 minutes goats' 
erythrocytes seen in gut 

14 

<J 

11 

ii 

11 

n 


Table VI.—Giving the results of feeding G. palpalis on defibrinated plasma through a membrane of rat skin 


No. 

Sex 

1 

Membrane 

Nature of fluid 

Result 

Remarb 

1 

<? 

Rat skin . 

70 % plasma and 30 % 
normal saline 

No visible distension 

After 15 minutes 

2 

9 ! 

>1 

11 

Slight distension 

11 

3 

<? 

n 

ii 

No visible distension 

11 

4 

<? 

ii 

11 

n 

11 

5 ! 

s 

n . 

11 

ii 

11 

6 i 

<? 

11 

11 

Almost complete 
distension 

11 

7 


11 

11 

No visible distension 

11 

8 

9 

” . 

n 

Complete distension 

11 


practically negligible. Two of eight flies, all of which had made 
repeated efforts to feed, succeeded in completely engorging 
themselves. In only one of the remaining six was slight distension 
noticeable. The results are given in Table VI. 


! 


l 












373 


From these experiments it is apparent that the most attractive 
element in the blood is the red corpuscle. In order to carry the 
matter further, the washed red cells of the goat were laked by the 
addition of three parts of distilled water to two parts of red cells. 
This solution was offered to three Glossina palpates, all of which 
quickly and completely engorged themselves, thus proving that the 
integrity of the red corpuscles is not an essential factor. An attempt 
was then made to separate the red cell stromata from those 
constituents of the corpuscle which are soluble in water. For this 
purpose sufficient sodium chloride was added to the laked red cell 
solution to render it isotonic, and the stromata were subsequently 
precipitated by prolonged centrifugalisation. Unfortunately, we 
did not succeed in obtaining complete separation of the stromata, as 
we had no high power centrifuge at our disposal, and owing to the 
minute size of the goat’s erythrocytes precipitation of the stromata 
is a matter of no small difficulty. Nevertheless, although complete 
separation was not obtained, a considerable fraction of the total 
amount of stromata was precipitated and a solution of goafs 
haemoglobin, fairly free from solid matter, was obtained. The 
resulting solution contained approximately as much haemoglobin as 
that found in a 40 per cent, suspension of goafs red cells. 

Four flies to which this solution was offered rapidly and 
completely distended themselves. The solution was then diluted 
with an equal volume of physiological saline. It was found that the 
six flies to which this solution was offered did not feed with the same 
avidity as did those which were offered the more concentrated 
solution; one became completely distended, three others partially, 
whilst in two no distension occurred. 

Having thus determined that Glossina palpates feeds readily on 
fresh haemoglobin solution, further experiments were performed with 
solutions made from the crystallised haemoglobin of commerce 
(Griiblefs dried haemoglobin). Solutions made from this prepara¬ 
tion are of a brownish-red colour, and give rather indistinct 
absorption bands of oxyhaemoglobin. The strength of the solution 
used was between 1 and 2 per cent. Four of the five flies to 
which this solution was offered became partly distended, whilst in 
the case of the other, although no distension was visible, the solution 
was seen in the gut on dissection. None were found to engorge 



374 


themselves completely in a manner comparable to that seen with 
haemoglobin solution made from fresh blood. Details are given in 
Table VII. 

Owing to the lack of suitable apparatus, we were unable to 
determine the result of offering to Glossina palpalis the washed 
stromata derived from red blood cells. 


Table VII.—Giving the results of feeding G. palpalis on haemoglobin solutions of various concentration 

through a rat skin membrane 


No. 

Sex 

Membrane 

Nature of fluid 

Result 

Remarks 

1 

? 

Rat skin . 

40 % # solution of goat 
haemoglobin in 0*9 % 
NaCl 

Complete distension 

Immediately 

2 

a 


n 

tt 

After 5 minutes 

3 

a 

n ••• ••• 

tt 

11 

Immediately 

4 

$ 

,, 

it 

tt 

it 

5 

? 

tt 

20 %• solution of goat 
haemoglobin in 0*9 % 
NaCl 

Partial distension ... 

After 15 minutes 

6 

$ 

n 

tt 

Slight distension 

it 

7 

$ 

tt 

n 

Almost complete 
distension 

tt 

8 

a 

11 

tt 

No visible distension 

tt 

9 

$ 

» ••• 

tt 

Slight distension 

tt 

10 

a 

11 

it 

No visible distension 

tt 

11 

$ 

it 

1 — 2% solution of dry 
crystalline haemoglobin 

Partial distension ... 

tt 

12 

a 

it 

n 

it 

tt 

13 

a 

1 

M 

tt 

tt 

tt 

After 15 minutes Hb. 
solution seen in anterior 
gut 

*4 

a 

It 

11 

No visible distension 

<5 

a 

n 

tt 

Slight distension 

After 15 minutes 


* By this is meant that the solutions contained respectively as much haemoglobin as do 40 and 20 P« 
cent, suspensions of goat red blood cells. 


Summarising the results of these experiments, we find that 
Glossina palpalis feeds with avidity, through rat’s skin, on fresh 
defibrinated blood and also on suspensions (50 to 5 per cent.) of 
washed red blood cells in normal saline, and on solutions containing 
as much dissolved haemoglobin as is present in 40 and 20 per cent, 
suspensions of red blood cells. Defibrinated plasma does not appear 
to have the same attraction for them, nor does sodium chloride 
solution alone, but the latter, containing a small proportion of the 
dried haemoglobin of commerce in solution is to a certain extent 












375 


taken up. Although these experiments are not quite so conclusive— 
especially as regards separation of the constituents of the red blood 
cells which are soluble and insoluble in water—as could be desired, 
nevertheless, we consider they suggest strongly that the element in 
the blood which is attractive to Glossina palpalis is that fraction of 
the erythrocyte which is soluble in water, most probably haemoglobin. 

Having completed these observations on blood, we decided to 
make use of the same technique with a view to determining whether 
Glossina palpalis will take up other solutions through fresh rat-skin. 
Reference has already been made to the fact that when sodium 
chloride solution was offered, although the flies repeatedly pierced 
the membrane, they did not appear to take up any of the fluid. At 
all events, no distension was noticed as a rule, although in one 
instance partial engorgement did apparently occur; it was by no 
means easy to state definitely whether or not any of the solution had 
been imbibed, as small quantities of a clear fluid cannot be 
recognised in the gut of the fly. In order to determine the point, 
the solution must be coloured with some dye that can be readily 
recognised. For this purpose methylene blue, neutral red and 
fuchsin were used. It was found that when physiological saline, 
containing these dyes in solution, was offered to Glossina palpalis 
small quantities were taken up by the flies in a proportion of the 
instances. Dissection showed that the intestinal tract was deeply 
stained, red or blue according to the dye used. In some cases the 
stain had spread to the salivary glands and to the fat bodies. In 
fact, so deeply were certain flies coloured that even before dissection 
the dye could be recognised through the integument of the abdomen. 
After obtaining this information various other solutions were offered 
to Glossina palpalis , details of which are given in Table VIII. It is 
interesting to note that three flies became completely engorged with 
0 9 per cent, sodium chloride solution to which had been added 
about 5 per cent, of cane-sugar and a little neutral red. Another fly 
became partially distended on a 25 per cent, solution of glycerine 
in water coloured with methylene blue. 

These observations demonstrate that under certain conditions 
Glossina palpalis will imbibe and even completely distend itself 
with fluids other than blood. Although in all the experiments just 
described the membrane used was the freshly removed skin of a rat 



376 


or rabbit, nevertheless we found that this tsetse will feed through 
other membranes, e.g., sheep’s bladder, peritoneal tissue and 
membrane composed of a thin sheet of rubber. As regards the last, 
one fly out of five completely distended itself with defibrinated blood 
through this membrane. Although the flies had not the slightest 

Table VIII.—Giving the result of feeding G. palpalii on various fluids through a membrane of skin 


No. 

Sex 

Membrane 

Nature of fluid 

Result 

Remarks 

1 

$ 

Fresh rabbit skin ... 

Methylene blue 

No visible distension, 

Anterior and mid-gut blue. 




solution* 

but abdomen of 
bluish tinge 

Posterior gut green. 
Salivary glands, mak 
pighian tubules and fat 
bodies blue 

2 

<? 

99 

it 

No visible distension 

99 

3 


Same rabbit skin 24 

>1 

99 

No staining seen on di»* 



hours old 



section 

4 

c? 

99 

>> 

99 

99 

5 

? 


t> 

99 

Proventriculus and anterior 






gut blue 

6 

cJ 



No visible distension, 

Whole intestine (except 
rectum), salivary glands, 





but abdomen of 





bluish tinge 

malpighian tubules and 
fat bodies deeply stained 

7 

? 

99 

Neutral red -i% solu- 

No visible distension 

Anterior gut red 




tion in water 



8 

cf 

99 


99 

99 

9 

9 

Fresh rat skin 

Fuchsin *1 % in *9% 

Slight distension 

Proventriculus and an tenor 




NaCl solution 

gut pink 

10 

? 

99 

t> 

99 

99 

11 

£ 

» 

Neutral red • 1 % solu¬ 

Considerable 

Whole gut (except rec¬ 




tion in water + sugar 

distension 

tum), salivary glands, 
malpighian tubules and 





Complete distension 

fat bodies deeply stained 

12 

9 

99 

>» 

! 99 

>3 

9 

»» 

99 

99 t 

i . 99 

14 

9 

99 

Methylene blue 

No distension 

Anterior and mid-gut deep 



solution + sugar 


blue 

15 

£ 

99 


99 

99 

16 

£ 

99 

99 

99 

Slight staining of anterior 






gut 

*7 

<3 

,, 

75% methylene blue 

99 

No staining 



solution + 25% 
glycerine 




18 

£ 

99 

99 

Partial distension ... 

Anterior and mid-gut and 






also salivary glands and 
fat bodies stained 

*9 

£ 

99 

t> 

99 

Anterior gut blue 


• The methylene blue solution used was the weak solution employed in Romanowsky's stain; its formula 
is given on page 369. 


difficulty in piercing the rubber, they did not seem able in the 
majority of cases to imbibe blood through it; possibly owing to the 
extremely elastic nature of the rubber the lumen of the proboscis was 
occluded, thus preventing the passage of fluid through it. 




377 


The practical point that we have to decide is, does Glossina in 
nature obtain food other than blood ? Although this question is not 
answered by the experiments described above, yet the results 
obtained are very suggestive. They show that under favourable 
conditions Glossina will take up solutions of vegetable origin, such 
as sugar and water. Apparently one essential is that the fluid 
should be presented enclosed by a membrane. The reason for this 


Tabls IX.—Giving the results of feeding G. palpdis on blood through membranes other than freshly-removed 

skin 


No. 

Sex 

Membrane 

Nature of fluid 

Result 

Remarks 

1 

a 

Rabbit skin removed 
24 hours previously 
and partially 
decomposed 

Defibrinated goat blood 

Complete distension 

Immediately 

2 

9 


11 

» 

»» 

3 

9 



Partial distension ... 

After 15 minutes 

4 

a 

Ox bladder. 


Complete distension 

Immediately 

5 

9 

n 


»» 

11 

6 

a 



Partial distension ... 

After 15 minutes 

7 

a 

»> 


Slight distension 

8 

a 

Peritoneal 
membrane, ox 


Partial distension ... 

»» 

9 

9 

» 

» 

Slight distension 

»> 

10 

a 

»» 


»* 


11 

9 

Thin rubber sheeting 

>1 


n 

12 

a 


n 

Complete distension 

!» 

*3 

9 

»> 


No visible distension 


*4 

a 

„ 



»> 

*5 

a 

it 


„ 


16 

9 



n 



is not that the flies require the fluid to be at a positive pressure before 
they can imbibe; it is possibly a mechanical difficulty, the fly being 
unable to take up fluid unless its proboscis is buried in the membrane 
or tissue. The experiments show, moreover, that even when the 
proboscis is inserted into a membrane, it does not necessarily follow 
that the fluid enclosed will be imbibed to any considerable extent. 
The results obtained prove most definitely that Glossina palpalis 
exhibits a preference for certain fluids, especially blood, red cells and 
haemoglobin. The physical character of the fluid does not, within 
limits, appear to matter. Defibrinated blood, a 50 per cent, 
suspension of red cells in saline, and a solution obtained by laking 
two parts of washed red cells with three of distilled water, are taken 







378 


up with equal readiness, whilst plasma and normal salt solution are 
only imbibed in small quantities as a rule. 

Turning to the question whether Glossina takes up vegetable 
juices in nature, we may state at once that we have not had the 
opportunity of deciding the point absolutely. Glossina of both 
sexes were frequently seen to plunge their proboscis into bananas, 
oranges and mangoes which had had their skin removed, and also 
into the skin itself. The insertion of the proboscis was repeated again 
and again, and the organ often remained buried in the fruit for a 
minute or more; the whole procedure certainly gave one the 
impression that the flies were endeavouring to obtain food from the 
fruit. Furthermore, we noticed that Glossina palpalis will pierce 
leaves placed on the surface of a fluid with the object apparently of 
imbibing the fluid below. 

Unfortunately, we had not time to extend our observations on 
this subject, but in view of the results already detailed, and of the 
observations of Carpenter who found vegetable tissue in a proportion 
of wild G . palpalis dissected by him, we consider that it is highly 
probable that tsetse do take up food of vegetable nature. The 
frequency with which a hungry fly will insert its proboscis, or at 
least attempt to do so, into practically any object presented—they 
can even be seen trying to pierce the glass vessel in which they are 
kept when no skin is in immediate contact with the glass—suggests 
that, in the absence of blood, they are on the look out for other food, 
for there is every reason to believe that Glossina recognises when it 
is in the presence of the skin of a living animal. 

Assuming Glossina palpalis cannot reproduce itself or live for 
any long period in the absence of blood, it is possible that food of 
a vegetable nature may suffice to enable it to exist in a locality which 
for a certain period in the year is denuded of vertebrates. 


CONCLUSIONS 

1. About eight per cent, of the wild G. palpalis in this district 
contain recognisable red blood cells—seven per cent, of mammalian 
origin and one per cent, nucleated red cells of unknown origin. 

2 . Seventy-two hours after G. palpalis had completely distended 
itself on rat’s blood recognisable red cells could no longer be found 



379 


in its intestine; after being fed on a fowl nucleated red blood cells 
could be recognised in 40 per cent, of cases at the end of a similar 
period. The flies were kept at a temperature of 8o°-86° F. 

3. Neither shed blood nor other fluid which is exposed (not 
covered by a membrane) can be imbibed by G. palpalis. 

4. G. palpalis can take up through a membrane of fresh skin 
not only blood and various dilutions of it with normal saline, but 
also suspensions of red blood cells in normal saline, and solutions 
of haemoglobin (both freshly made from red blood cells, and the 
dried crystalline preparation of commerce) in distilled water. 

5. Fluids other than blood such as solutions of sugar, sodium 
chloride, and glycerine, in water containing a small quantity of a 
dye (methylene blue, neutral red or fuchsin) are also taken up 
through a membrane of fresh skin by G. palpalis, but not so quickly 
or so readily as is blood. 

6. G. palpalis exhibits a definite selective taste for the various 
fluids presented to it under the membrane; blood, red cells, and 
haemoglobin solution being much preferred. The attractive element 
in the blood is the fraction of the red cells soluble in water, probably 
haemoglobin. 

7. G. palpalis which had been starved for a day or two can often 
be seen to insert the proboscis repeatedly into oranges, bananas or 
other fruits which may be offered them. 

8. We are of opinion that G. palpalis in nature may under 
certain conditions take up fluid other than blood. 



380 


REFERENCES 

Carpenter, G. D. H. (1912). Progress Report on Investigations into the Bionomics of 
Glossina palpalis , July 27, 1910, to August 5, 1911. Report of the Sleeping Sickness 
Commission of tbe Royal Society , No. XII, pp. 79-111. 

- (1913). Second Report on the Bionomics of Glossina fuscipes {palpalis ) of Uganda. 

Reports of tbe Sleeping Sickness Commission of tbe Royal Society , No. XIV, pp. 1-37. 

Decent (1909). Sleeping Sickness Bulletin , Vol. I, p. 471. 

Lloyd, Ll. (1912). Notes on Glossina morsitans in Northern Rhodesia. Bull. Ent. Res., 
Vol. Ill, pp. 95-96. 

- C 1 913)- Glossina morsitans in the laboratory. Ann. *Irop. Med. & Parasit ., Vol. VII, 

pp. 285-292. 

Maugham (1911). Sleeping Sickness Bulletin , Vol. Ill, p. 271. 

Rodhain, J., Pons, C., Vandenbranden, J., and Bequaekt, J. (1912). Contribution au 
Mecanitme de la Transmission des Trypanosomes par les Glossines. Arcb. f. Scbiffs- u. 
Trop. Hyg.y B. 16, S. 732-739. 

Stuhlmann, F. (1907). Beit rage zur Kenntnis der Tsetsefliege. Arb. a. d. Kaiserlicben 
Gesundbeitsamte , B. 26, S. 301-383. 

Taute, M. (1912). Expcrimentelle Studien iiber die Beziehungen der Glossina morsitans 
zur SchUfkrankheit. Zeitscbr. f. Hygiene , B. 72, S. 316-320. 



THE CULTIVATION OF THE LEPROSY 

BACILLUS 

HENRY FRASER, M.D. Aberd. 

DIRECTOR, INSTITUTE FOR MEDICAL RESEARCH, FEDERATED MALAY STATES 

AND 

W. FLETCHER, M.D. Cantab. 

PATHOLOGIST, INSTITUTE FOR MEDICAL RESEARCH, FEDERATED MALAY STATES, 

KUALA LUMPUR 

(Received for publication 16 ]une y 1915) 

In an article entitled ‘ Leprosy: a perspective of the results of 
experimental study of the disease/ by Bayon, and published in the 
Annals of Tropical Medicine , Vol. IX, the statement is made on 
page 29 that ‘ Fraser and Fletcher discarded all diphtheroids because 
of their ubiquity/ It is unfortunate that he has quoted our work 
incorrectly. In the Lancet , Vol II, 1913, we published an article 
entitled ‘The Bacillus leprae : has it been cultivated?* and on 
page 920, under the head of contaminating micro-organisms, we 
state that * In common with other workers we have isolated 
diphtheroid organisms, but these are ubiquitous and demand no 
special consideration.* At the time that article was written, our 
investigations had led us to form the opinion that the diphtheroids 
were contaminators, and had no genetic relationship with the leprosy 
bacillus; further investigations carried out during the past two years 
have confirmed that opinion. 

If we had confined our observations to the results of one or two 
experiments, as has been done by so many workers, we might, 
perhaps, like Bayon, have formed the opinion that the diphtheroids 
were related to the leprosy bacillus, but to do so we should have had 
to reconcile or, since that was impossible, to ignore some remarkable 
discrepancies. 

It seems a comparatively simple operation to reflect the skin from 
a leprous nodule and to excise a portion of the subcutaneous tissue 
free from contamination, but only those who have performed the 
operation a sufficient number of times can be aware of the pitfalls. 

Cultures of diphtheroids were obtained only in our early experi¬ 
ments ; in these a nodule of tissue rich in leprosy bacilli was excised, 
and from twenty to thirty tubes of culture media were inoculated 



382 


with portions of it. On one tube, perhaps two tubes, a culture of a 
diphtheroid would be obtained. As each tube was inoculated with 
an enormous number of leprosy bacilli from the same nodule,- it is 
impossible to believe that only in one or two parts of that nodule 
was there an acid-fast bacillus capable of proliferation under 
saprophytic conditions into a diphtheroid. If every tube inoculated 
with portions of the nodule had developed a culture of a diphtheroid 
the opinion formed might have been quite different, but the sporadic 
occurrence of a diphtheroid is quite in accordance with our results 
obtained in the isolation of diphtheroids from other parts of the 
body, and we believed the correct interpretation of these experiments 
to be that the nodule was contaminated, or had become contaminated 
from the skin, with one or two diphtheroid bacilli which proliferated 
on the culture media. 

In that belief we extended our investigations and improved our 
technique, avoiding the use of all disinfectants for the skin. We 
were then able to excise leprous nodules free from contamination, 
and from which a culture of no organism could be obtained on any 
medium. 

We have now excised material from fifty-two non-ulcerating 
nodular cases of leprosy, a number far in excess of that recorded by 
any other worker, and have employed the media of every claimant 
to success, but have not substantiated the claims of any of them. 

Our work has been carried out over a period of three and a half 
years The only conclusions, which that work permits, are that the 
leprosy bacillus has not been cultivated and that the diphtheroids 
and other organisms are merely contaminators. 

Elsewhere we (1914) have dealt with Kedrowsky's culture of an 
acid-fast bacillus which Bayon claims to be the leprosy bacillus. 
Our experiments have led us to conclude that there is no evidence 
that the acid-fast bacillus of Kedrowsky is the leprosy bacillus. 


REFERENCES 

Bayon, H. (1915). Leprosy: a perspective of the results of experimental study of the 
disease. Annals Trop. Med. & Parasitol ., IX, pp. 1-90. 

Fxasxr, H., and Fletchki, W. (1913). The Bacillus leprae: has it been cultivated? 
Lancet , Sept. 27, 1913, pp. 918-921. 

- - ( I 9 , 4)- Fourteenth Annual Report of the Institute for Medical 

Research, Kuala Lumpur, Federated Malay States. 



383 


THE RESERVOIR OF THE HUMAN 
TRYPANOSOME IN SIERRA LEONE 

[Being the Third. Report of the Thirty-second Expedition of the 
Liverpool School of Tropical Medicine, 1914-1915.] 

BY 

WARRINGTON YORKE 

AND 

B. BLACKLOCK 

(Received for publication 24 June , 1915) 

No time was spent in examining the native population for the 
incidence of trypanosomiasis, but two cases of sleeping sickness— 
a woman and her child—were brought to our notice by Drs. Young 
and Butler of the West African Medical Service. These cases which 
were extremely chronic were discovered accidentally, one or two 
trypanosomes being seen during the examination of the blood for 
malarial parasites. The patients lived in Lumley, a native village 
situated at the junction of the Cape Lighthouse Peninsula with the 
mainland, about a mile and a half from Hill Station and four miles 
from Freetown. Trypanosomes were first seen in the mother 
(Catherine Macauley) about the beginning of 1914. She received a 
single treatment with salvarsan, and subsequently returned to 
Lumley. When seen by us on December 7th, 1914, she was in good 
condition. According to Dr. Young, no definite change had 
occurred during the year. The only symptoms were headache and 
a feeling of weakness. She had one or two slightly enlarged glands 
in the posterior triangles. Several careful examinations of the 
peripheral blood and gland juice failed to reveal trypanosomes. On 
the day she was first seen by us (December 7th) two rats were 
inoculated with blood and one with gland juice. After 28 days one 
of the two rats inoculated with the blood was found to be infected 
with trypanosomes. The other two rats were kept under observa¬ 
tion until they died, on the 59th and 67th days respectively; 
trypanosomes were never found in them. There was no change in 



the condition of the patient when she was last seen by us on 
February 24th, 1915. On December 16th, 1914, she brought her 
son, aged 8, to the hospital suffering with fever. On examination 
of a stained preparation of his blood by Dr. Butler, malarial 
parasites were found and also a single trypanosome. On 
December 19th the boy was seen by us. He complained of pain 
over the spleen, which was enlarged, reaching to within one inch of 
the umbilicus. There were a few shotty glands in the neck, axillae 
and groins. Malaria parasites were still present in the blood, but 
no trypanosomes were seen. Two rats were inoculated with 1 c.c. 
of the peripheral blood. Neither of these animals became infected; 
one died on the 52nd day afterwards, and the other was still alive 
on the 90th day when we left the Colony. The blood of this boy 
was examined by us on several other occasions, but trypanosomes 
were not seen. 

The history of these two cases shows that sleeping sickness in 
Sierra Leone is exceedingly chronic, and very difficult to recognise. 
Trypanosomes can be found only occasionally in the peripheral 
blood; gland puncture of the mother was negative, whilst the boy 
had no glands which were puncturable. 

The question whether there is any reservoir of the human 
trypanosome in Sierra Leone other than man is of importance, and 
is one which we decided to investigate. By the term Reservoir we 
mean the source from which Glossina derives its infection. 
Kinghom and Yorke (1912) have shown that the chief reservoir of 
the human trypanosome of South Central Africa (T. rhodesiense ) 
is the antelope and not man. The antelope differs from man in 
being tolerant of the infection; unlike man, in whom the disease 
runs an acute course, it is able to harbour T. rhodesiense in its 
blood for long periods without exhibiting signs of the disease. Both 
for this reason and because the antelope exists in enormous numbers 
in many parts of South Central Africa, coinciding in its distribution 
with the ubiquitous G. morsitans —the fly responsible for the 
transmission of sleeping sickness in this region—it constitutes a 
much greater and more reliable reservoir of the human trypanosome 
than does man. In Sierra Leone, however, we have to deal with 
quite a different state of affairs; sleeping sickness in this part of 
Africa is an extremely chronic infection. In striking contrast to 



3»S 


cases infected with T. rhodesiense , those infected with the human 
trypanosome of the West Coast may live for many months, or even 
years, without exhibiting any serious indication of the disease. For 
this reason, therefore, man in Sierra Leone is a more constant and 
dependable reservoir of the virus of sleeping sickness than he is in 
Rhodesia or Nyasaland. Moreover, in the Colony of Sierra Leone, 
and in most portions of the Protectorate, large game is compara¬ 
tively rare. Again, the most prevalent tsetse-fly in Sierra Leone 
is G. palpalis ; owing to its predilection for water-courses its distri¬ 
bution is limited much more closely to the haunts of man than is 
that of the ubiquitous G. morsitans. In Sierra Leone, therefore, we 
must conclude that of large game and man, the latter constitutes by 
far the more important reservoir of the human trypanosome. 

There is, however, another possible reservoir of the human 
trypanosome which must not escape attention, that is domestic 
stock. The Sleeping Sickness Commission of the Royal Society 
(1910) examined seventeen cattle in Uganda, and found one cow 
infected with T. gambiense. This element does not exist in 
Rhodesia or Nyasaland, as quite apart from T. rhodesiense cattle 
are unable to thrive to any extent in the presence of large numbers 
of G. morsitans. Although cattle are not bred in most portions of 
Sierra Leone, yet they are imported from French Senegal in large 
numbers, and gradually find their way down to Freetown for 
slaughter. 

Both in the Protectorate and in the Colony the blood of a 
number of domestic animals—143 animals, 7 goats, 7 sheep and 
10 dogs—was examined. As will be seen in a subsequent paper, 
the trypanosomes most commonly found were T. vivax and 
T. congolense. One ox, however, was found to be infected with a 
trypanosome which we are unable to distinguish from that infecting 
man. The ox in question was one of a herd of 90 Government 
cattle in a Warri at Batkanu. All the animals in this herd had 
originally come from French territory, but they had been in the 
Protectorate for various periods before being bought by the Govern¬ 
ment. The animal in which this trypanosome was found appeared 
to be in perfect health; direct examination of its peripheral blood 
was negative. The trypanosome was discovered fortuitously owing 
to the fact that this ox happened to be one of nine chosen at random 



from the herd for the purpose of having their blood inoculated into 
rats. Trypanosomes were seen in the peripheral blood of the rat 
fourteen days after inoculation. The parasite was a polymorphic 
trypanosome, it did not exhibit posterior nuclear forms, and was 
indistinguishable morphologically from the trypanosome isolated 
from the human case of trypanosomiasis. 

During the last six months the strain has been maintained by 
passage through rats. The course of the infection in these animals 
is exceedingly chronic, and certain rats failed to become infected. 
For purpose of comparison, the result of inoculation of rats and 
guinea-pigs with this strain and with that obtained from man are 
given in tabular form. 

There is a striking similarity in the pathogenicity of the two 
strains in rats. The virulence of both is but slight, and is in marked 
contrast to that of the other common polymorphic trypanosomes— 
T. pecaudi or T. rhodesiense vel ugandae (T. brucei , Uganda). On 
consulting the results of inoculation of rats with T. gambiense direct 
from man (Yorke, 1910) one cannot fail to observe their striking 
similarity to those set forth in the above table. Macfie (1914) 
records that he failed to infect rats with the Nigerian strain of the 
human trypanosome (T. niget tense). A guinea-pig infected with 
T. nigeriense was, however, sent by Macfie to us at Runcorn, and 
from it we succeeded without much difficulty in infecting a number 
of rats. The course of the infection in these animals was 
exceedingly chronic often ending in recovery; some failed to become 
infected. 

We are of opinion, therefore, that this polymorphic trypanosome 
from the ox is identical with that infecting man in West Africa, 
in other words, that it is T. gambiense . 

The discovery of the human trypanosome in the ox is important. 
It shows that domestic stock may serve as a reservoir of 
T. gambiense. Before we can form any opinion as to the relative 
importance of this reservoir we must have information on the 
following two points : — 

(1) Do the animals harbour the parasite in their blood for long 
periods without exhibiting signs of disease, or does the infection 
run an acute course? 

(2) What percentage of the animals is infected ? 



3«7 


Table I.—Giving results of inoculation of rats with the human strain of T. gambiense 


No. of Animal 

Animal from which 
inoculated 

Incubation 

in 

days 

Length 
of life 
in days 

Remarks 

Rat la . 

Patient’s blood 

28 

■56 


» ib . 

33 j» 

— 


Did not become infected 

» io* . 

Rat la . 

8 

88 


» iob . 

» « . 

13 

— 

Alive on 171st day. 
Trypanosomes last seen 
on 50th day 

» 37 * . 

33 * oa . 

4 

29 


» 37 ^ . 

>» ioa . 

— 

— 

Did not become infected 

33 5 Ia . 

n loa . 

— 

- 

33 >» 

>1 5*b . 

5j i° a . 

— 

— 

» i» 


Table II.—Giving results of inoculation of rats and guinea-pigs with the ox strain 
of T. gambieiue 


No. of Animal 

Animal from which 
inoculated 

Incubation 

in 

in days 

Length 
of life 
in days 

Remarks 

Rat 24 . 

Ox VIII . 

14 

40 


» 3 i . 

Rat 24 . 

7 

— 

Alive on 160th day. 
Trypanosomes last seen 
on 24th day 

» 47 a . 

» M . 

10 

39 


» 47 b . 

3) H . 

10 

— 

Alive on 139th day. 
Trypanosomes last seen 
on 67th day 

33 49 ® . 

3, 3 » . 

— 

- 

Did not become infected 

t> 49 b . 

33 3 i . 

— 

— 

13 11 

33 50 . 

33 47 b . 

10 

63 


Guinea-pig 52 

33 5 ° . 

— 

— 

Did not become infected 

Rat 53* . 

33 50 . 

18 

- 

Alive on 46th day 

». 53 b . 

33 5 ° . 

18 

— 

33 33 

Guinea-pig 54 

33 53 a . 

— 

— 

Did not become infected 

55 . 

33 53 a . 

*3 

— 

Alive on 31st day 



























388 


As regards the first point, the ox from which the trypanosome 
was obtained appeared in perfect health, and a report received four 
months later states that there is still no indication of disease. 
Regarding the second point, we have as yet no evidence to show 
what proportion of the cattle in Sierra Leone are infected with 
T. gambiense. If, as appears probable, cattle prove to be tolerant 
of this parasite and can harbour it in their blood for long periods 
without detriment to health, it will be no easy matter to determine 
how many animals are infected. As already mentioned, the 
parasite was discovered largely by chance; although a direct 
examination of the peripheral blood of each of the 90 animals in 
the Warri was made, subinoculation into rats was done in nine 
instances only. Furthermore, it must be borne in mind that rats 
not infrequently fail to become infected after inoculation of blood 
containing T. gambiense. It will, therefore, probably be at least as 
difficult to determine the percentage of cattle infected as it is to 
ascertain the number of infected human beings. 

Before leaving the subject, it is interesting to refer to the 
observations recorded by Macfie. This investigator holds that in 
the Eket district man does not constitute a reservoir of sleeping 
sickness, and that the reservoir is still unknown. The evidence on 
which he bases this assumption is, in our opinion, unconvincing. 
He writes 4 The extreme rarity or complete absence of trypanosomes 
from the peripheral blood of all the human cases examined, the 
rarity of the parasites even in the gland juice, and the difficulty 
experienced in infecting animals by inoculation, suggest that it 
must be a very exceptional occurrence for a tsetse-fly to become 

infected by feeding on these cases.It seems, therefore, that 

the human infections must be dependent on some other cycle of 
development, including an insect and some so far unidentified 
animal host, the reservoir of the disease. The ordinary development 
of the trypanosome may take place in these two hosts. The insect, 
infected from the animal host may, however, be capable of infecting 
human beings; but the disease may be so modified in them that they 
are incapable of handing on the infection any further/ This 
argument appears fallacious. In view of the fact that trypanosomes 
were not found in the peripheral blood they were doubtless scanty, 
but it does not by any means follow that they were absent. Nor 



389 


does the failure to infect animals by inoculation of blood materially 
assist the argument, since Macfie himself found what has frequently 
been pointed out by previous workers that rats and guinea-pigs 
often fail to become infected after the injection of blood known to 
contain T. gambiense. Macfie records that all his attempts to 
infect rats with blood containing the trypanosomes were unsuccessful; 
further, of seven guinea-pigs inoculated with gland juice actually 
containing trypanosomes only one became infected. 

This comparative insusceptibility of rats and guinea-pigs to 
human trypanosomes of the West Coast shows that we cannot safely 
regard a negative result obtained by inoculation as evidence that 
trypanosomes were not present in the. blood inoculated. It is, 
therefore, inadmissible to argue, as does Macfie, from the negative 
results of microscopic examination of the blood and of inoculation 
of guinea-pigs and rats that G. falpalis cannot become infected from 
human beings. In 5 c.c. of the blood of one of these patients there 
may be sufficient trypanosomes to infect many flies, and yet 
insufficient to infect a particular guinea-pig or rat. 

Briefly, Macfie’s view appears to be this. Man is not a reservoir 
of the virus of sleeping sickness because Glossina cannot become 
infected from him. He postulates an unknown vertebrate reservoir 
of the virus and an insect host, in which the ordinary development 
of trypanosomes takes place. The insect may infect man but cannot 
become infected from him, but only from the hypothetical vertebrate 
reservoir. 

For the reasons given above, we consider that Macfie has failed 
to show that Glossina cannot become infected from man. In our 
opinion, there is no evidence which would lead us to believe that man 
does not constitute a most important reservoir of the human 
trypanosome of West Africa. Nevertheless, the fact that the human 
trypanosome has been discovered in an ox indicates that domestic 
stock also forms a reservoir, the extent of which we are at present 
unable to estimate, but the existence of which must be recognised 
when prophylactic measures are contemplated. 

CONCLUSIONS 

1. The human trypanosome, T. gambiense , has been found in 
an ox in Sierra Leone. 



390 


2. It is impossible at present to form any conclusion of the 
extent to which domestic stock may harbour this parasite. In all 
probability, it will prove to be at least as difficult to recognise the 
infection in cattle as it is in man. 

3. The existence in domestic stock of a potential reservoir of 
T. gambiense will have to be taken into account when prophylactic 
measures are contemplated. 


REFERENCES 

Bruce, Hamerton, Bateman, Mackie and Lady Bruce (1910). Experiments to ascertain if 
cattle act as a reservoir of the virus of Sleeping Sickness ( Trypanosoma gambiense). Free. 
Roy. Soc. B. VoL 82, pp. 480-484. 

Kinghorn, A. and Yorke, W. (1912). On the Transmission of Human Trypanosomes by 
Glossina mortitans, Westw.; and on the Occurrence of Human Trypanosomes in Game. 
Annals Trap. Med. and Parasitol ., VI, pp. 1-24. 

Macfie, J. W. S. (1914). Sleeping Sickness in the Eket District of Nigeria. Annals Trap. 
Med. and Parasitol ., VIII, pp. 379-438, with 5 plates. 1 map. 

Yorke, W. (1910). On the Pathogenicity of a Trypanosome (T. rbodesiense , Stephens and 
Fantham) from a case of Sleeping Sickness contracted in Rhodesia. Annals Trap. Med • 
and Parasitol ., IV, pp. 351-368. 



39* 


SPIROCHAETA BRONCHIALIS , 
Castellani, 1907, TOGETHER WITH 
REMARKS ON THE SPIROCHAETES OF 
THE HUMAN MOUTH 

BY 

H. B. FANTHAM, M.A. Cantab., D.Sc. Lond. 

LECTURER ON PARASITOLOGY, LIVERPOOL SCHOOL OP TROPICAL MEDICINE. 

[Being the First Report of the Thirty-first Expedition of the 
Liverpool School of Tropical Medicine , to Khartoum , 1913] 

{Received for publication 1 ]une> 1915) 

Plate XXXIV 

CONTENTS 


I. Introduction. 391 

II. Material and Methods . 392 

III. Movements . 394 

IV. General Morphology . 395 

V. Morphological Variation in Spirocbaeta broncbialu . 396 

VI. The Granule Phase or Spirocbaeta broncbialu . 399 

VII. Some Further Remarks on the Granule Phase or Spirochaetes. 401 

VIII. The Spirochaetes or the Human Mouth . 402 

IX. The Spirochaetes or the Human Mouth contrasted with S. broncbialu ... 404 

X. Mode or Intection. 404 

XI. General Remarks . 405 

XII. Summary and Conclusions. 408 

Reverences . 409 

Explanation or Plate . 412 


I. INTRODUCTION 

During the summer of 1913, I had the good fortune to work in 
the Wellcome Tropical Research Laboratories, Khartoum, having 
been sent on expedition by the Liverpool School of Tropical 
Medicine. While there, thanks to the kindness of the Government 
of the Anglo-Egyptian Sudan and of the Director, Dr. A. J. 
Chalmers, and Staff of the Wellcome Laboratories, I was able to 


















39 2 


undertake researches on human and avian spirochaetes, directing 
especial attention to bronchial spirochaetosis in man. In this 
memoir an account is given of some of my researches on the causal 
agent of the latter disease. 

II. MATERIAL AND METHODS 

The material was chiefly obtained from cases of human bronchial 
spirochaetosis occurring in Khartoum, Omdurman and Kodok. 
Many of the cases were chronic. They included officers’ servants, 
warders, native policemen, soldiers and household servants. A few 
cases among Europeans suffering from bronchitic symptoms were 
studied, and some of them were found to be fresh cases of undoubted 
spirochaetosis. Among the latter were two cases of almost experi¬ 
mental infection. The total number of cases studied was twenty. 

I was fortunate in being able to obtain material from nearly all 
the cases reported by Drs. Chalmers and O’Farrell (1913). I was 
also able to examine preparations sent by Dr. J. A. Taylor from 
Entebbe, Uganda, to whom my best thanks are due. In fact, 
Dr. Taylor sent me his first preparations for my examination and 
opinion at the end of May, 1913, just before I left England for the 
Sudan. Spirochaeta bronchialis was also studied in monkeys 
experimentally infected in Khartoum. 

It is with much pleasure that I acknowledge the kindness of 
Dr. Chalmers and Captain O’Farrell, R.A.M.C.; Major Forrest, 
R.A.M.C.; Major Carroll, R.A.M.C.; Dr. Christopherson; 
Dr. Atkey; Dr. Crispin; Captain Stirling, R.A.M.C., and 
Captain Buist, R.A.M.C., in providing me with material for research. 
I am especially indebted to the gentlemen first-named for much help 
while in the Sudan. 

The subject is one of great difficulty, and because of this 
difficulty and the pressure of other work the publication of this paper 
has been delayed. Also, even now, it is not so fully illustrated 
as I should have wished. Many more drawings (totalling over 500) 
have been prepared, but owing to the war these cannot be reproduced 
without further delay. Hence a selection of what appear to be 
types of the spirochaetes only are shown on the accompanying 
plate (XXXIV), the reproduction of which by half-tone process has 
left something to be desired. 



393 


Also, owing to the lack of precise knowledge of spirochaetes 
found in the human mouth and respiratory passages, attention has 
been paid to the comparative morphology of these organisms. I 
have compared the spirochaetes of the mouth found in natives of 
the Sudan with those found in similar situations in people living in 
England, especially in cases of pyorrhea alveolaris and of ‘ gum¬ 
boils,* obtained in Liverpool and in Cambridge. 

In the Sudan the procedure followed for obtaining material was 
briefly as follows:—The teeth were examined for caries, as were 
also the buccal, tonsillary and pharyngeal secretions, with a view 
to finding spirochaetes. The patient washed his mouth with clean 
water, and samples of spirochaetes in mouth sputum might also be 
thus obtained. The throat was then gargled, and usually samples 
of throat sputum were examined. After this careful washing of the 
mouth and gargling of the throat, expectoration from the deeper 
bronchial regions was collected in a sterile petri-dish, and in this 
Spirochaeta bronchialis occurred. The sputum thus collected was 
examined by dark-ground illumination, and films were made with 
the aid of a sterile platinum loop for immediate fixation and 
staining. 

Various methods of fixing and staining were tried. Osmic 
vapour followed by absolute alcohol, Schaudinn's fluid (sublimate- 
alcohol-acetic) and Bouin*s fluid (picro-formol-alcohol-acetic) were 
used for fixing wet preparations; Bouin*s fluid was found most 
useful. The solutions of Giemsa and Leishman, carbol thionin and 
gentian violet, were tried, as well as various haematoxylins, such as 
those of Delafield, Ehrlich and Bohmer. In the hot dry atmosphere 
of Khartoum, freshly prepared Bohmer’s haematoxylin was found 
to be most useful, and less complicated than iron-haematoxylin, 
though this Was also used. Staining overnight with dilute 
solutions was found to be more effective in the case of this 
fragile spirochaete than quick staining with stronger solutions. 
Unfortunately, gentian violet, so useful for spirochaetes in general, 
was found in Khartoum to fade quickly, but was very useful when 
studying mouth spirochaetes in England. Naturally, in the fixation 
of a spirally wound, flexible organism, some flattening and 
straightening of the coils is inevitable, the fixed forms thus being 
less graceful structures than the living ones. 



394 

III. MOVEMENTS 

In fresh sputum 5 . brottchialis moves very quickly. Its move¬ 
ment, like that of other spirochaetes, is difficult to analyse. The 
organism moves forward while turning on its long axis. However, 
as I have previously described (1907, 1908), such movements may 
be divided into two components, namely, an undulatory flexion of 
the body mainly for progression, and a corkscrew or helicoid move¬ 
ment of the body as a whole, due to the winding of the membrane 
or crista. This latter organella is very narrow and thin, and is a 
lateral outgrowth of the periplast. It is only seen with difficulty in 
S. brotichialis , and not in some specimens, as it is sometimes so 
contracted as to appear to be absent. It does not markedly 
undulate, and should not be called an undulating membrane in 
spirochaetes, as I pointed out in 1907-8, for it is not directly 
comparable with the undulating membrane of a trypanosome. It 
can be seen in some cases in life by the use of the paraboloid 
condenser. In stained specimens it sometimes appears as a curved 
line lying along the body when the organism has been Axed during 
rapid movement (PI. XXXIV, Ags. 2, 22), or as a slightly wavy 
lateral outgrowth when the organism was killed while moving slowly, 
the membrane being somewhat relaxed (Ags. 10, 16, 36). 

The undulatory flexion is responsible for the rippling motion of 
spirochaetes and the waves seen passing down the body of these 
organisms. 

Slowly moving spirochaetes have few undulations of relatively 
large amplitudes along their bodies, while quickly moving forms 
have more numerous small undulations (Ags. 19, 23, 28), characters 
which I also described in 1907-8. Hence the number of coils or 
waves in a spirochaete is more an index of its rate of motion—as well 
as of its thickness,which is also a factor—than one of differentiation 
between various forms. Thus, thin forms are more easily thrown 
into waves than thick forms, and so the number of waves in a 
spirochaete cannot be used as a mark of species differentiation as 
has been attempted by some workers in the past. In spirochaetes 
exhibiting polymorphism, thin forms represent young individuals, 
while thick ones are adult forms, as was indicated by me in 1909. 

Jerking movements may occur, but not frequently. Spirochaetes 
are isopolar, and can easily and very quickly return on their own 
path. Other remarkable appearances are produced by rapid move- 



395 


ments in a relatively limited area. Thus, the spirochaete may curve 
each of its ends inwards in a watchspring-like manner and after a 
time straighten itself, and then re-form a watchspring coil at either 
end in the opposite direction. Waves continue to pass along the 
organism during the movements of the body as a whole. At 
other times, though more rarely, the organisms intercoil their ends 
and seem almost to tie themselves in knots, only to extend a few 
seconds later and swim away in a different direction. The subject 
of the movements of spirochaetes was fully discussed by me in June 
and August, 1907, and January, 1908, when dealing with 
S. balbianii and S. anodontae , and the same remarks apply to 
S. bronchialis and the spirochaetes of the mouth. There was little 
evidence of division seen in life. 

IV. GENERAL MORPHOLOGY 

The morphology of spirochaetes is a subject of great difficulty 
and of no little controversy. In this paper I shall, without prejudice, 
adopt the generic name Spirochaeta, as I cannot at present accept the 
unconfirmed researches of Zuelzer (1911) on that debatable organism 
known as S. plicatilis. As a result of Zuelzer’s researches some 
workers have proposed to create a number of genera such as 
Cristispira (for the forms found in molluscs), Spiroschaudinnia or 
Borrelia (for blood-inhabiting forms), Treponema , and the like, but 
without any clearly and generally accepted generic differences. The 
divergent descriptions of S. plicatilis given by different workers may 
be clearly seen and contrasted from the account given in Bosanquet's 
book on Spirochaetes. The so-called axial fibre of Zuelzer is 
acknowledged to be homologous with the membrane or crista of 
molluscan spirochaetes. Subsequent researches may lead to a 
solution of these difficulties. 

Spirochaeta bronchialis is a delicate organism which frequently 
stains with difficulty. It exhibits morphological variation (figs. 1-39), 
due to growth and division. There are long forms (figs. 37-39) 
and short forms (figs. 1-6); also thick ones (figs. 7, 38) and thin 
ones (figs. 28, 29, 33), and those of intermediate length (figs. 7-33) 
and breadth (figs. 25, 34, 35). No detailed account of the morphology 
of the parasite has yet been published. 

The cytoplasm of S. bronchialis is almost homogeneous and 



39 $ 


shows no vacuoles. In spite of the minuteness and fragility of the 
organism, chromatin bars can be seen at intervals along the parasite 
in some of the specimens (figs. 14, 24, 29, 36-39). In stained 
preparations the observance of these minute bands or rodlets is some¬ 
times aided by the use of stereoscopic eyepieces. When examined 
fresh these chromatin granules appear as retractile spots, as 
viewed under dark-ground illumination using a paraboloid 
condenser. On the other hand, many spirochaetes only show the 
chromatin dots or granules with difficulty at certain stages of their 
life-cycle, and so may appear more or less homogeneous (figs. 19, 
28, 34)- 

The presence of chromatin bars or rodlets at short intervals 
along the spirochaete in a stained preparation (figs. 24, 29, 32) gives 
rise to a so-called alveolar or chambered appearance, which has been 
much emphasised by Gross (1910) and those who have followed him. 
There is little or nothing new in Gross’s observations, except the 
terms used to express his interpretations. However, his views have 
been so dogmatically and even polemically asserted that work 
previous to that of Gross has tended to be overlooked, although it 
was performed in many cases with due regard to careful cytological 
technique. The accentuation of differences of interpretation is 
merely indicative of partisanship, and does not conduce to the 
progress of knowledge. 

The ends of S. bronchialis are tapering. Observed separately 
they may appear to be pointed (figs. 26, 33) or somewhat rounded 
(fig. 7) in stained preparations. Sometimes one end appears rather 
more tapering than the other (figs. 31, 39), a feature that is 
explained by extending attenuation at a previous division. So-called 
flagella are also thus explicable. These variations in the character 
of the ends of S. bronchialis also occur in other spirochaetes, such as 
S. balbianii , and were described by me at some length in 1909. 


V. MORPHOLOGICAL VARIATION IN SPIROCHAETA BRONCHIALIS 

Spirochaeta bronchialis exhibits considerable polymorphism, 
resulting from the processes of growth and division. This range of 
variation is responsible probably for the varying dimensions of the 
organism given by different workers. The difference of size 



397 


exhibited by S. bronchialis can only be realised by examination and 
measurement of a large number of specimens derived from a series 
of cases. Measurements of spirochaetes from one or two cases only 
may be very fallacious, since the organisms observed may be all 
practically at the same stage of development. 

By measuring a moderately large number (300) of bronchial spiro¬ 
chaetes, the discrepancies in length may be explained; I find that the 
length ranges from 5 p to 27 p. The size of a number of them 
centres around 15 p, while many of the others are about 8 p 
long. Castellani and Chalmers (1913) in their 'Manual of Tropical 
Medicine/ state on p. 1283 that one form of S. bronchialis is from 
15 P to 39 At, while on p. 402 the commonest form is said to be 
7 p to 15 p long. Thus my measurements give a lower limit for both 
minimum and maximum dimensions as stated by them, while a 
number of the spirochaetes that I have measured are about 15 p, 

Macfie (1915) states that his smallest form was 6 p long, his 
largest 13 p , and that the average length of his specimens was 8 p to 
9 p . When examining my series of slides, I found one—taken from 
a chronic case on a particular day—on which the spirochaetes seemed 
remarkably uniform. A number was drawn with the aid of a camera 
lucida and measured (figs. 7, 12, see also 13). It was found that 
almost everyone of them was from 8 p to 9 p long. Their measure¬ 
ments coincided almost exactly with those given by Macfie, and 
had this slide been the only one, erroneous conclusions would have 
resulted. Measurements of the spirochaetes from the same case on 
the days preceding and on subsequent dates showed that the range 
of length of 5 . bronchialis was considerably greater than was shown 
on that one occasion. As Macfie's dimensions were derived from 
two cases only, it was likely that the majority of the spirochaetes 
were at the same stage of development, and thus had attained about 
the same size. Many bronchial spirochaetes from Khartoum were 
either from 14 p to 16 p long, or 7 p to 9 p long. Definite 
evidence of transverse division in members of the former group 
has been obtained, and it is very likely that Macfie’s forms 
measuring 8/1 to 9 m so originated. Chamberlain (191 1) having 
found bronchial spirochaetes in two typhoid patients in the 
Philippines, states that their average length is 15 p. Some of 
the spirochaetes that I examined in the Sudan measured about 



398 


15 ft long, but much depended on the age of the parasites and on 
the condition of the patient. For instance, the predominant type 
of S. bronchialis found in several chronic cases was an organism with 
two tapering ends, one sometimes slightly more rounded than the 
other, with a length of 13 ft to 16 ft (figs. 34, 35). 

Dr. Taylor kindly sent me some slides of S. bronchialis from 
Uganda. I have drawn and measured spirochaetes from each of the 
slides he sent me, and can state that not only can I recognise the 
four types mentioned by him, but that the dimensions which I have 
obtained from his sketches agree in the main with mine (figs. 40-45). 
The range of length obtained from my measurements is from 5*5/1 
to 19 5 ft. Dr. Taylor, in his Report for 1913, differentiated four 
groups of spirochaetes, which I have determined from his figures to 
measure from 6 fit to 9 fit, 6 fit to 8 fit, 15 fit to 18 ft and 9 ft to 11 ft, 
respectively. The first three were considered by him to be bronchial 
spirochaetes, the last occurred in the mouths of healthy persons. 
While the spirochaetes were thus grouped by Dr. Taylor, he recog¬ 
nised the existence of possible transitional forms and that one group 
gradually merged into another. 

The length of S. bronchialis is often a factor of its age. Forms 
recently developed by the elongation of granules or coccoid bodies, 
naturally, are short (fig. 1-6). Small forms also arise by transverse 
division of older, longer ones. The nutrient medium in which the 
spirochaetes occur also seems to react upon their morphology. 
Organisms surrounded by thick stringy mucus may become stouter 
than those in more fluid sputum, which offers less resistance to their 
passage through it. Spirochaetes found in a denser medium tend 
to be both shorter and broader than those in a more fluid one in 
many cases, though no general statement can be made regarding this 
factor. The degree of relaxation obtaining in the organism when it 
was killed is also a factor in producing differences in the appearance 
of the coils of the organism, and consequently in its dimensions. 

Polymorphism or morphological variation of S. bronchialis occurs 
not only with respect to length, but also in connection with breadth 
and the character of the ends of the organism. The breadth of the 
spirochaetes shows some variation, but it is less marked than the 
variations in length. Occasionally a spirochaete broader than the 
average is found (fig. 38). The breadth is not easily determined, 
but varies from 0*2 ft to 0*6 ft . 



399 


The ends of the spirochaetes show considerable variation. The 
degree of tapering manifested by them depends in part on the motion 
of the parent organism during division, as mentioned previously. 
Spirochaetes in which both ends taper, but wherein one is somewhat 
more pointed than the other, are fairly common (figs. 27, 34, 37, 39, 
46). Others show the ends equally pointed (figs. 8, 10, 22, 26); yet 
others have relatively rounded ends after some tapering of the body 
has occurred (figs. 7, 31, 47, 48). Extreme tapering of one or both 
ends of the spirochaete have led to the erroneous interpretation of 
flagella in the case of other spirochaetes, such as S. dentium . 
Occasionally such an interpretation might be made in specimens of 
5 . bronchialis , but examination of many of the organisms, both 
living and in stained preparations, confirms the fact that true flagella 
do not occur in the organism. 

A membrane or 4 crista* is present in S. bronchialis (figs. 10, 16), 
but is not always easy of detection. This difficulty of demonstrating 
the membrane would, perhaps, cause some workers to assign the 
organism to the genus Treponema , in which hitherto a membrane has 
not been satisfactorily demonstrated. Such classification, however, 
would be fallacious, inasmuch as the coils of a Treponema are said 
to be pre-formed while those of a spirochaete are not, but are capable 
of variation according to the rapidity or slowness of motion and to 
the density of the medium traversed. 

VI. THE GRANULE PHASE OF SPIROCHAETA BRONCHIAUS 

The formation of coccoid bodies or granules in Spirochaeta 
bronchialis has been studied in fresh preparations, using both 
ordinary illumination and the paraboloid condenser, as well as in 
stained preparations. As the organisms are small, there is some¬ 
times some slight difficulty in studying the process of formation of 
these minute reproductive bodies, though it is relatively easy to 
observe in larger, somewhat stouter specimens about 11 n to 16 fi long 
and about 0*3 fi broad examined at the right stage of development. 
Further, there seems to be a periodicity in the formation of granules 
in S. bronchialis , but the exact period has not yet been determined. 

The process of formation as observed in life is as follows: — 
The cytoplasm at first is very finely granular, in fact, almost 
homogeneous. The chromatin bars appear as minute refractile 



400 


masses. A concentration of some of the cytoplasm occurs around each 
chromatin rodlet. These small concentrations gradually become 
oval, the outer cytoplasmic layer differentiates as a thin coat, and 
ultimately a series of coccoid bodies or granules is formed 
(figs. 46-49), lying usually transversely or slightly obliquely within 
the periplast sheath. Sometimes the coccoid bodies are set at 
liberty by a rupture appearing at one end of the spirochaete (fig. 50), 
at other times several ruptures, or disintegration of the sheath can be 
observed. A few empty sheaths have been found both in fresh and 
in stained preparations. Stained specimens show a series of darker 
lozenge-like coccoid bodies alternating with relatively clear, pale 
staining areas (figs. 47-49). 

The formation of coccoid bodies takes place both in free 
spirochaetes and in those which have penetrated the delicate cells 
lining the air passages (fig. 52). Sometimes two or more spirochaetes 
have been found within a mononuclear cell. They may be in the 
ordinary trophic phase, or may be in process of formation of 
coccoid bodies. When within a cell the coccoid bodies often 
seem to be liberated by the disintegration of the periplast. Groups 
of coccoid bodies still retaining the outline of the spirochaete from 
which they originated (fig. 52) are of fairly frequent occurrence. 
When the coccoid bodies are released by a terminal rupture of the 
parent, they tend to form irregular clumps. The elongation of the 
granules and the emergence of very small spirochaetes from the 
groups of granules (fig. 51) have been observed in life. It is very 
probable that there is a definite period in the life of a spirochaete at 
which there is a marked differentiation of coccoid bodies. It must 
also be borne in mind that coccoid bodies may be present when 
spirochaetes as such cannot be detected. 

In staining reactions, size and morphology, the coccoid bodies 
are different from any bacteria which may be present in the 
surrounding medium. They stain less darkly than cocci, and are 
smaller (fig. 52). Spirochaetal coccoid bodies have no capsule such 
as occurs commonly in bacteria. 



4oi 


VII. SOME FURTHER REMARKS ON THE GRANULE 
STAGE OF SPIROCHAETES 

In continuance of the remarks which I made on the subject of the 
granule stage of spirochaetes in these Annals (1914), Vol. VIII, 
pp. 471-484, I should like to draw attention to the following further 
evidence. 

1. Noguchi, in his address before the Royal Society of 
Medicine, London, on October 20th, 1913, stated that he ‘was able 
to demonstrate .... granules in the pure cultures of Treponema 
pallidum . This phenomenon, however insignificant it may appear 
in itself, was destined to furnish a key to one of the most disputed. 
problems of the past fifty years—namely, the problem of so-called 
parasyphilis, since it was this very idea that prompted me to 
undertake to search for Treponema pallidum in one form or another 
in the brains of general paralytics and in the spinal cord from cases 
of tabes dorsalis.' And again, ‘ I was led by the observation that 
Treponema pallidum sometimes assumes a granular form in cultures 
to re-study sections of paretic brains stained for th z pallidum? 

2. Another interesting remark regarding the granule stage of 
spirochaetes may be found in Sir Patrick Manson’s well-known book 
on Tropical Diseases. In the last (fifth) edition, published in 1914, 
on page 225, regarding the etiology of relapsing fever, he writes: 
‘Obermeier and von Jaksch describe certain refractile bodies present 
in the blood during the fever intermissions. The latter author says 
that he has observed the development of these bodies into short rods, 
from which the typical spirochaetes are eventually evolved.’ These 
remarks have also appeared in earlier editions of the book. 

3. Dutton and Todd, writing in 1905 concerning some of their 
experiments on the nature of human tick fever in the Congo Free 
State, record that: ‘ In some preparations of stomach or malpighian 
tubules [of spirochaete-infected ticks] no parasites were at first seen; 
but if a little human serum, taken from one who had never had tick 
fever, were added, in from 8 to 24 hours the preparations became 
fairly crowded with spirochaetes.* These observations may be 
explained by spirochaetes passing through a granule phase. . 



402 


VIII. THE SPIROCHAETES OF THE HUMAN MOUTH 

Two species of spirochaetes were recorded as occurring in the 
human mouth about forty years ago. These are S. buccalis, 
Steinberg (often ascribed to Cohn, 1875), and 5 . dentium , Miller 
(often attributed to Koch, 1877). The former is said to be longer 
and thicker than the latter. The smaller spirochaete is also the 
more flexible. According to Hartmann and Miihlens (1906), 
S. dentium in culture measures 4 /* to 12 /* in length by 0*3/1 to 
0*6 fi in breadth, or less if stains other than Loffler's are used, while 

S. buccalis is 12 p to 20 /* in length by 0*5 fi to 1 /* in breadth. 
Hoffmann and Prowazek (1906) published microphotographs of 
specimens of 5 . dentium about 5 /* to 7*5 /* long, and of 
5 . buccalis about 8*5 n to 14*5 ft long. Intermediate types also 
occur, as was shown by these authors, such forms being named 
Treponema intermedium by Dobell (1912). Hartmann and 
Miihlens did not see much internal structure in S. dentium , but in 
5. buccalis they record the presence of an ‘ undulating * membrane in 
some specimens. Miihlens (1907) figured stained specimens of 
5 . buccalis and S. dentium in which chromatin coloured granules 
were distributed along the bodies of the organisms. His figures of 
5. dentium, as measured by me, are about 4 fi to 7/1 long and of 
5 . buccalis about 15 fi to 23 /*. 

Noguchi (1912) has succeeded in cultivating a number of species 
of spirochaetes from the human mouth. He places these in the genus 
Treponema , making three new species, namely, T. macrodentium , 

T. microdentium and T . mucosum , but they cannot be easily 
distinguished morphologically. The last-named was obtained from 
cases of pyorrhea alveolaris. They all differ from S. buccalis. 

In the course of my researches I have observed the parasites 
ascribed to Cohn and to Koch, these being the two common spiro¬ 
chaetes seen in the mouths of natives of the Sudan and of Europeans 
in England, as well as the forms described and cultivated by later 
investigators. Some of the mouth spirochaetes are not very active, 
but there is marked corkscrew and boring movement, and they are 
flexible. Tangles or tomenta of these mouih spirochaetes are common. 
Internal structure is seen with some difficulty, but in some specimens 
it can be determined, and chromatin granules are then observed. 



4°3 

S. dentium (h gs. 53-57) has tapering ends, and varies in length from 
4 fi to 10 fi, as found by measuring 40 specimens from some of 
my own preparations. S. dentium is so small that few details of its 
structure can be determined, and in consequence, it might be placed 
by some authorities in the genus Treponema. S. buccaiis 
(figs. 58-68) has somewhat rounded or bluntly acuminate ends, and 
varies in length from 9 fi to 22 /1, as determined from 110 specimens 
measured. A slight membrane or crest may sometimes be observed 
in the latter species (figs. 60, 65). Intermediate forms were found, 
which might be considered to connect the two species. 

The spirochaetes of the mouth take up stains well and with 
relative ease. Intracellular stages, if they occur at all, are 
uncommon. Multiplication by binary fission has also been observed. 
Coccoid bodies or granule stages of the mouth spirochaetes are 
formed, but appear to be relatively few in number. 

No marked morphological difference has been observed between 
the spirochaetes occurring in the mouths of Europeans in England 
(four cases examined), and in natives of the Sudan (eight cases 
examined). However, it may be of interest to note that in the Sudan 
the predominant spirochaete in the mouths of the natives whom 
I examined was S. buccaiis , S. dentium being relatively uncommon. 

Spirochaetes that were either 5 . dentium or closely allied thereto 
were found in the mucus from the pharyngeal and laryngeal regions 
of a normal monkey in the Sudan. 

A spirochaete of the human throat, often associated with 
fusiform bacilli, may be mentioned here. Spirochaeta vincenti 
occurs in the throat in certain conditions, such as Vincent’s angina. 
It has recently been described by Drs. J. G. and D. Thomson (1914). 
It is said to be an elongate spirochaete, with a flexible, irregularly 
coiled body, tapering at both ends. The spirochaete, according to 
the present state of our knowledge, does not seem to be identical with 
5 . bronchialiSy differing from it morphologically and in the situations 
in which it is found. S. vincenti was named by Blanchard in 1906, but 
as no description of the spirochaete was given at the time of naming, 
the species name is probably not valid. Little is known of the 
range of morphological variation of S. vincenti. I have measured 
those drawn by Thomson and find that they are from 10 fi to 18/1 
long, while the few figured by Miihlens (1907) I find to measure 



9 Hj io (i and 23/1. Casteilani and Chalmers state that it is 12 ^ 
to 25 n long. It is possible that this species will have to be merged 
into another one, such as 5 . bronchialis, when it has been more fully 
investigated. 

IX. THE SPIROCHAETES OF THE HUMAN MOUTH CONTRASTED 
WITH S. BRONCHIAUS 

There are general differences between these two groups of 
spirochaetes, and these differences may be set forth with a view to 
aiding those workers in the tropics who may meet with cases of 
bronchial spirochaetosis. However, the said differences must not be 
emphasised too much or asserted dogmatically, as further work may 
lead to modifications. 

The bronchial spirochaetes appear to be more active than those 
of the mouth. 

5 . bronchialis dies very rapidly outside the respiratory tract. 
This has also been noticed by Chalmers and OTarrell and by 
Taylor. Oral spirochaetes, on the other hand, can live for some 
hours outside the human mouth. 

Bronchial spirochaetes, as a rule, stain with difficulty and not so 
easily as those obtained from the mouth. The former are much more 
fragile and slender than 5. buccalis. 

Coccoid bodies are frequently produced by 5 . bronchialis, and 
are probably the cross-infective stage of the parasite. Coccoid 
bodies appear to be less frequently formed by oral spirochaetes. 

Intracellular stages of S. bronchialis are occasionally seen. No 
such stages have been observed in the cases of spirochaetes from the 
mouth, which I have examined. 

Tangles are not usually formed by 5 . bronchialis, but are 
commonly produced by numbers of spirochaetes aggregating in 
clusters in the oral cavity. 

X. MODE OF INFECTION 

The situation in which Spirochaeta bronchialis is found largely 
precludes its dissemination direct from man to man by the aid of 
insects, though they may serve as indirect transmitters of the 
parasite. 



It is highly probable that man himself acts as the reservoir of the 
virus, and that direct contact of infected with healthy persons is 
responsible for the spread of bronchial spirochaetosis. The spray 
exhaled with the expired air appears to be contaminated with the 
resistant coccoid bodies produced by S. bronchialis. Such fine spray 
inhaled by an uninfected person whose bodily resistance is somewhat 
lowered—as after a chill—results in an attack of bronchial 
spirochaetosis. It is possible that spirochaetes as such might be 
exhaled in the spray, but as has been pointed out they are fragile 
and quickly die outside the body. It is also possible that the 
inhalation of dust contaminated with dry sputum from patients 
suffering from bronchial spirochaetosis may excite an attack, while 
the nasal secretions passed on to linen and allowed to dry, the linen 
then being packed indiscriminately with other soiled clothing, may 
serve to transmit the parasite to new hosts. 

The agency of insects in spreading bronchial spirochaetosis is 
somewhat remote, but it must be remembered that certain flies are 
partial to sputum, which they not only ingest but which may soil 
their bodies and feet. While the frail, motile spirochaetes 
are not likely to be carried as such in or on the bodies of flies, it is 
quite possible that the more resistant coccoid bodies may be carried 
about by flies, and deposited by them either directly on the lips or 
nose of man, or in foods such as milk, whence they may reach the 
pharynx and perhaps pass downwards into the trachea and bronchi. 

However, the direct contamination of healthy persons by infected 
ones is at present much the most likely means of dissemination of 
5 . bronchialis . 


XI. GENERAL REMARKS 

Certain points of interest connected with Spirochaeta bronchialis 
and bronchial spirochaetosis may now be considered. 

That Spirochaeta bronchialis is disease producing is undoubted. 
The organisms have been found in cases of chest complaints, 
especially those with bronchitic symptoms, where no other cause for 
the affection was assignable. S. bronchialis was frequently the only 
organism present in the cases in the Sudan examined by me. 
Examination of a series of preparations from several chronic cases, 



406 


each considered separately, has shown that, on most of the occasions 
on which sputum was examined, bacteria of any kind were either few 
or absent. Sometimes organisms identical morphologically with 
Diplococcus ( Micrococcus ) catarrhalis were found on a single 
preparation of a series extending over a number of days, but this was 
uncommon. Occasionally bacteria were found in smears of mouth 
spirochaetes. On still rarer occasions cells from the lungs were 
observed in which endocellular spirochaetes and coccoid bodies 
related thereto were present in company with diplococci (fig. 52). 
The contrast between the coccoid bodies and the bacteria, both in 
size, morphology and staining reactions, was always noticeable. In 
one case only were pneumococci found associated with Spirochaeta 
bronchialis , the patient being a European soldier who might have 
become infected with spirochaetes after a chill had favoured 
pneumonic symptoms. No mycological agent has been found in any 
of my preparations. Chalmers and O’Farrell also have had a similar 
experience, and they state that the fungi responsible for broncho- 
moniliasis, broncho-nocardiasis, broncho-aspergillosis, broncho- 
penicilliosis, broncho-mucor-mycosis, broncho-sporotrichosis, have 
not been found by them 4 in the sputum of their cases of bronchial 
spirochaetosis either by direct examination or by cultivation.* 
Further, Chalmers and O’Farrell succeeded in experimentally 
inducing bronchial spirochaetosis in a monkey, the parasites and 
symptoms closely resembling those in man (cf. figs. 19, 49). 

It is also of interest to note that Chalmers and O’Farrell, and 
Taylor, both writing in 1913, arrived at the same conclusions 
regarding bronchial spirochaetosis in Khartoum and Uganda 
respectively. In each district S. bronchialis is held to be the cause 
of the haemorrhagic bronchitis noticed. Also each of the above 
workers state that there is an abundance of S. bronchialis during the 
attack, that the decrease in numbers synchronises with the abatement 
of symptoms, and that the spirochaetes disappear from the sputum 
entirely or are found with great difficulty after a few days. All 
these facts have a distinct significance when considering the etiology 
of the disease. 

5 . bronchialis I find to be morphologically different from the 
spirochaetes of the oral cavity, as mentioned previously. It is an 
entity in itself, and the malady caused by it also appears to be 



distinctive. Its pathogenic action also seems to be unlike that of the 
debatable S. vincenti of Vincent’s angina, in which a throat 
membrane may at times be produced. 

S. bronchialis has a somewhat extensive though scattered 
distribution. Bronchial spirochaetosis has been reported from 
Europe, the cases being discussed by Chalmers and O’Farrell. It 
has also been recorded from Ceylon, India, the Philippines, and the 
West Indies. In East Africa the disease has been reported from the 
Anglo-Egyptian Sudan and Uganda. In West Africa the disease 
has been recently notified from various parts of the Gold Coast 
Colony. Spirochaeta bronchialis has thus been reported from four 
continents and from different districts in each. This scattered but 
wide distribution of the parasite suggests that it may be a very 
general but easily overlooked organism, of far more frequent 
occurrence than is usually supposed. 

For the benefit of medical men who are interested in the subject 
and to whom Chalmers and O’Farrell’s paper is not available, the 
treatment recommended by them is here quoted : — 

1 The first essential is rest in bed, good food, and ventilation. 
The second is arsenic in some form, preferably associated with 
glycerophosphates. These may be given by the mouth with excellent 
results, or intramuscularly as an injection of: — 

Sodium cinnamate . 0 05 grm. 

Sodium cacodylate . o’10 ,, 

Sodium glycerophosphate . 0*10 ,, ’ 

Taylor, in Uganda, prescribes ‘ Arsenious acid by the mouth in 
increasing doses.’ 

At the time when these treatments were recommended (end of 
1913) they had been employed in relatively few cases, but had given 
satisfactory results. 

Chalmers and O’Farrell, and Taylor also discuss the 
co-existence of bronchial spirochaetosis with other chest diseases, 
such as lobar pneumonia, details of which should be sought in the 
original papers. Bronchial spirochaetosis may be suspected in 
atypical cases of pneumonia and bronchitis, and may be mistaken 
for incipient phthisis. 



XII. SUMMARY AND CONCLUSIONS 


1. Spirochaeta bronchialis is an organism presenting marked 
polymorphism, a feature that has only been determined by the 
examination of numerous preparations from various patients. It 
produces bronchial affections in the Sudan and in other parts of the 
world. 

2. 5 . bronchialis , as investigated in the Anglo-Egyptian Sudan, 
varies in length from 5 fi to 27 fi 9 and its breadth is about 0*2 p to 
0 6 fi . These variations are due to the processes of growth and 
division. Many of the parasites measure either 14 fi to 16 fi long, 
or 7 fi to 9 fi , the latter resulting from transverse division of the 
former. The ends show much variation in form, but approach the 
acuminate type on the whole. The discrepancies in dimensions 
given by the very few previous workers on S. bronchialis are the 
result of the measurement of a limited number of parasites. All such 
sizes can be found on some occasion during the progress of the 
disease, when a larger number of spirochaetes is examined. 

3. The movements of S. bronchialis are active, but of relatively 
short duration, when it is removed from the body. The number of 
coils of the spirochaetes is rather an index of its rapidity of motion 
than a fixed characteristic of the species. 

4. The motile phase of S. bronchialis is succeeded by one of 
granule formation, the granules or coccoid bodies serving as a resting 
stage from which new spirochaetes are produced. The formation of 
coccoid bodies and the reproduction of spirochaetes from them can 
be observed in life. 

5. S. bronchialis is a species distinct from the spirochaetes 
occurring in the mouth. It differs from them in morphology, 
pathogenicity and in staining reactions. It is not a developmental 
form of any bacterium, and is an entity in itself. 

6. The passage from man to man is effected most probably by 
means of spirochaetes, and especially coccoid bodies, that leave the 
body in the spray with expired air and by way of the nasal 
secretions. Owing to the fragility and short life of S. bronchialis 
extracorporeally, the resistant coccoid bodies in air, dried sputum 
and dust, and possibly also on the bodies of flies and other insects, 
are probably instrumental in inducing attacks of bronchial spiro- 



4°9 

chaetosis in human beings, especially those having a lowered bodily 
resistance, such as after a chill. 

7. Spirochaetes of the type of S. dentium , measuring 4 fi to 10 fi 
in length, have been found in mouths of natives of the Sudan and in 
those of English people in England. Also S. buccalls , measuring 
9 fi to 22 fiy has been obtained from the same sources. 

8. S. bronchialis will probably prove to be of more frequent 
occurrence than is known at present. 


REFERENCES 

Relating to Bronchial Spirocbaetosis 

Castellani, A., and Chalmers, A. ]. (1913). Manual of Tropical Medicine, 2nd edition, 
pp. 402 and 1283. 

Chalmers, A. J., and O’Farrell, W. R. (1913). Bronchial Spirochaetosis. Journ. Trop. 
Med. and Hyg ., Vol. XVI, pp. 329-334. 

Chamberlain, W. P. (1911)* The Occurrence in the Philippines of Associated Spirochaetae 
and Fusiform Bacilli in Ulcers of the Throat (Vincent’s Angina), of the Mouth, and 
of the Skin, and in Lesions of the Lungs (Bronchial Spirochaetosis). Philippine Journ. 
Science , B, Vol. VI, pp. 489-498. 2 Pis. 

- (1914)* Spirochaetae and Fusiform Bacilli in various lesions in the Philippines. Amer. 

Journ. Trop. Dm., Vol. II, pp. 246-255. 1 PI. 

Harper, F. S. (1914). Bronchial Spirochaetosis. Journ. Trop. Med. and Hyg Vol. XVII, 
p. 194. 

Macfie, J. W. S. (1915). Bronchial Spirochaetosis. Journ. Trap. Med. and Hyg., Vol. 
XVIII, pp. 63-65. 

Taylor, J. A. (1914). Bronchial Spirochaetosis in Uganda, with pneumonic symptoms. 
Annals Trop. Med. and Parasitcl ., Vol. VIII, pp. 13-18. 

- (1914)* Bronchial Spirochaetosis in Uganda. Annual Med. and Sanit. Rept., Uganda 

Protectorate , for 1913, pp. 80-86. 


Relating to Spirochaetes of the Human Mouth 

Hartmann, M., and MOhlens, P. (1906). Untersuchungen iiber Bau und Entwicklung der 
Zahnspirochaten. Zeitscbr. f. Hyg. und Infektionskrankb., Bd. LV, pp. 92-112. 2 Pis. 

Hoffmann, E., and Prowazek, S. von (1906). Untersuchungen iiber die Balanitis- und 
Mundspirochaten. Centralbl. /. Bakt ., 1 Abt., Orig., Bd. XLI, pp. 741-744, 817-821. 
1 PI. 

MOhlens, P. (1907). Vergleichende Spirochatenstudien. Zeitscbr. f. Hyg. und Infektions¬ 
krankb., Bd. LVII, pp. 405-416. 3 Pis. 



410 


Noguchi, H. (1912). Cultural Studies on Mouth Spirochaetae £ Treponema microdentivm 
and macrodentium). Journ. Exper. Med., Vol. XV, pp. 81-89. 5 Pis. 

- (1912). Treponema mucosum (new species), a mucin-producing spirochaeta from 

pyorrhea alveolaris, grown in pure culture. Journ. Exper. Med., Vol. XVI, pp. 194-198. 
1 PI. 

Thomson, J. G., and Thomson, D. (1914). Some Researches on Spirochaetes occurring in 
the Alimentary Tract of Man and some of the Lower Animals. [Marcus Beck Lab. 
Repts.—No. 3.] Proe. Roy. Soc. Med., Vol. VII, Pt. 1, pp. 47-70. 2 Pis. 


Relating to Spirochaetes in General 


Blanchard, R. (1906). Spirilles, Spirochetes et autres microorganismes A corps spirals. 
Arch, de Parasitologic, Vol. X, pp. 129-149. 

Bosanquct, W. C. (1911). Spirochaetes. Pp. 152. W. Saunders Company, Philadelphia 
and London. 

Dobell, C. C. (1912). Researches on the Spirochacts and related Organisms. Arch. /. 
Protistenkunde , Bd. XXVI, pp. 117-240. 5 Pis. 

Dutton, J. E., and Todd, J. L. (1905). The Nature of Human Tick-Fever in the Eastern 
Part of the Congo Free State. Liverpool Scb. Trap. Med., Memoir XVII. See p. 17. 

Fa nth am, H. B. (1907). Spirochaeta {Trypanosoma) balbianii (Certes), its Movements, 
Structure, and Affinities j and on the Occurrence of Spirochaeta anodontae (Keysselitz) 
in the British Mussel, Anodonta cygnea. (Preliminary Account.) Ann. & Mag. Nat. 
Hist., ser. 7, Vol. XIX, pp. 493-501. 

- (1907). The Movements of Spirochaetes, as seen in S. balbianii and S. anodontae. 

Brit . Assoc. Rept., 1907 (Leicester), pp. 554-355. 

- (1908). Spirochaeta (Trypanosoma) balbianii (Certes) and Spirochaeta anodontae 

(Keysselitz) : their Movements, Structure, and Affinities. Quart. Journ. Microsc. Sci., 
Vol. LII, pp. 1-73. 3 Pis. 

-(1908). The Spirochaetes : A review of some border-line organisms between animals 

and plants. Science Progress, Vol. Ill, No. 9, pp. 148-162. 4 Text-figs, 

- (1909). The Spirochaetes found in the Crystalline Style of Tapes aureus: a Study 

in Morphological Variation. Parasitology, Vol. II, pp. 392-408. 1 PI. 

-(1911). Some Researches on the Life-Cycle of Spirochaetes. Annals Trop. Med. 

and Parasitol., Vol. V, pp. 479-496. 6 Text-figs. 

- (1914). The Granule Phase of Spirochaetes. Annals Trop. Med. and Parasitol., 

Vol. VIII, pp. 471-484. 2 Text-figs. 

Fantham, H. B., and Porter, A. (1909). The Modes of Division of Spirochaeta recurrentis 
and 5 . duttoni as observed in the Living Organisms. Proc. Roy. Soc., B, Vol. LXXXI, 

pp. 500-505* 



Gross, J. (1910). Cristispira nov. gen. Ein Beitrag zur Spirochatenfrage. Mittb. a. d. zool. 
Station zu Neapel , Bd. XX, pp. 41-93. 1 PI. 

- (1912). Ueber Systematik, Struktur und Fortpflanzung der Spironemacea. Centralbl. 

/. Baku , I Abt., Orig., Vol. LXV, pp. 83-98. 

Noguchi, H. (1913). On some of the Recent Advances in the Field of Microbiology; with 
Demonstrations of the Pure Cultures of various Spirochaetes, of the Viruses of Rabies 
and Poliomyelitis, and of Treponema pallidum in the Brains of General Paralytics. 
[Occasional Lectures.] Proc. Roy. Soc. Med. y Vol. VII, Pt. 1, pp. 3-30. (See pp. 5, 19.) 


Zuelzer, M. (1911). Ueber Spirocbaeta plicatilis Ehrbg. und dcren Verwandtschafts- 
beziehungen. Arch. f. Protistenkunde , Bd. XXIV, pp. 1-59. 4 Pis. 



412 


EXPLANATION OF PLATE XXXIV 

All figures were outlined with an Abb6-Zeiss camera lucida, using a 

Zeiss 2 mm. apochromatic objective and compensating ocular 18. The 

magnification is approximately 2,600 diameters. 

Figs. 1-39. Spirochaeta bronchialis from the Anglo-Egyptian Sudan. Trophic 
stage. 

Figs. 1-6. Small, young forms of S. bronchialis. 

Figs. 1,3. Spirochaetes with both ends slightly differing in form. 

Figs. 2, 4. Forms with both ends alike. In fig. 2 the edge of the membrane 
shows as a line on the body. 

Figs. 6-13. S. bronchialis . Growth forms showing variation in thickness and 
length. 

Fig. 10. Spirochaete with relaxed membrane. 

Fig. 13. Spirochaete drawn from preparation obtained from Kodok, Sudan. 

Figs. 14-24. Longer forms. 

Fig. 16. Form with slight membrane. 

Figs. 19, 23. Forms with many small coils, fixed when in rapid motion. 

Fig. 19 drawn from specimen from an experimentally infected 
monkey. 

Figs. 14, 20. S. bronchialis from a case at Kodok. 

Fig. 22. The edge of the membrane (crista) shows as a wavy line on the 
body. 

Figs. 25-35. Some of the commoner (‘average’) types of S. bronchialis , 
showing variation in the number of coils. 

Figs. 36-39. Some of the largest forms of 5 . bronchialis encountered. 

Fig. 36. Spirochaete with slight membrane. 

Fig. 37. Form with one end more pointed than the other. 

Fig. 38. Very broad form. 

Fig. 39. Long form with slight membrane. 

Figs. 40-45. Spirochaeta bronchialis from Uganda. 

Figs. 40-44. Trophic forms showing the same appearances and structure 
as the average types of S. bronchialis occurring in the Sudan. 

Fig. 45. Parasite showing the formation of coccoid bodies. 

Figs. 46-52. The granule phase of S. bronchialis from cases in the Sudan. 

Figs. 46-49. Free spirochaetes containing coccoid bodies. Fig. 49 drawn 
from a preparation obtained from an experimentally infected 
monkey. 

Fig. 50. Coccoid bodies in process of liberation at one pole of the spirochaete. 

Fig. 51. Cluster of small coccoid bodies with young spirochaetes emerging 
from the group. 

Fig. 52. Part of cell from the air passages, containing two spirochaetes, a 
group of coccoid bodies retaining the arrangement of the spiro¬ 
chaete from which they were liberated, and two diplococci. 

Figs. 53-57. Spirochaeta dentium from mouths of Europeans and natives of the 
Sudan. 

Figs. 53, 54. S. dentium , from natives of the Sudan. 

Figs. 55, 56. 5 . dentium , from Case 1, Cambridge. 

Fig. 57. S. dentium , from Case 2, Cambridge. 

Figs. 58-68. Spirochaeta buccalis from mouths of Europeans and natives of 
the Sudan. 

Figs. 58, 62, 63, 65. S. buccalis , from Case I, Cambridge. Fig. 65 shows a 
membrane or crista. 

Fig. 59- S. buccalis , from Case 2, Cambridge. 

Fig. 64. S . buccalis , from Case 1, Liverpool. 

Figs. 60, 61, 66-68. S. buccalis , from natives of the Sudan. In fig. 60 a 
slight membrane is indicated. 



Annals Trap. MeJ. & Parasite!., Pol. IX 


PLATE XXXIP 



SPIROCHAETA BRONCHIALIS, Figs. 1—52 
SPIROCHAETES FROM HUMAN MOUTH, Figs. 53—68 




4'3 


NOTES ON CERTAIN ANIMAL 
PARASITES OF DOMESTIC STOCK IN 
SIERRA LEONE 

• 

\Being the Fourth Report of the Thirty-second. Expedition of the 
Liverpool School of Tropical Medicine, 1914-15.] 

BY 

WARRINGTON YORKE 

AND 

B. BLACKLOCK 


(Received for publication 29 June, 1915) 


SERIOUS EPIDEMICS OF DISEASE AMONGST CATTLE IN 

SIERRA LEONE 

From time to time heavy losses have been experienced by cattle 
owners in Sierra Leone owing to epidemics, during which large 
numbers of cattle die. 

In December, 1914, we were requested by the Administration to 
investigate the nature of a serious epidemic which had broken out 
amongst cattle in the northern part of the country. The cattle in 
question had been brought down originally from French Senegal; 
they were practically all bullocks, owing to restrictions respecting 
the export of cows put into force by the French Administration. 

We proceeded to Batkanu, the capital of the Karene district, but 
were informed on our arrival that the epidemic had ceased and fhat 
the animals were no longer dying. We found 90 cattle in the 
Government Warri, all of which appeared to be in a healthy 
condition. Most of the animals were in the Karene district during 
the epidemic, and had remained apparently uninfected; a few, 
however, had been ill, and seemed to have recovered. Whether 



these animals had really suffered from the infection which had 
resulted in the death of large numbers of others is uncertain, but 
Dr. Clearkin, the medical officer in charge of the district, was of the 
opinion that this was the case. We were informed that the epidemics 
occurred towards the end of the rainy season, and that they 
coincided with the appearance of large numbers of a fly {Stomoxys 
nigra), which was held responsible. This hypothesis is in 
accordance with the history given by the natives; those questioned 
by us stated that the disease appeared every five or six years, 
during the rainy season, and that it was due to the bites of the fly 
(Stomoxys). It was believed that the infection was contracted by 
the animals whilst being brought down from French country, where 
cattle breeding is carried on extensively. 

The course of the disease was rapid, the symptoms being fever, 
staring coat, emaciation, and death in a month or six weeks. 

Fresh and stained preparations of blood of each of the 90 cattle 
in the Warn were examined, and rats were inoculated with blood 
from every tenth animal. As a result of this examination four 
cattle were found to be infected with trypanosomes; two with 
T r vivax r, one with T . congolense and one with T. gambiense. In 
two of the animals Piroplasma bigeminum was found, and in about 
30 per cent, a parasite belonging to the genus Theileria. Two 
healthy oxen were inoculated with blood from each of the animals 
suffering from infection with T . vivax . The animals inoculated 
from the first ox remained healthy, and trypanosomes were never 
seen in their blood; those inoculated from the second, which also 
harboured Piroplasma bigeminum in its blood, became infected both 
with T. vivax and with Piroplasma bigeminum , but exhibited no 
signs of disease, and were alive and in good condition when we left 
the colony three months later. 

In view of the fact that we did not see these animals until the 
epidemic had ceased, it was impossible for us to make any post¬ 
mortem examination, and there was no material at our disposal 
upon which a definite conclusion could be reached as to the cause of 
the great mortality which had occurred. Although the examination 
of blood proved that the cattle were infected with several protozoa, 
any one of which is potentially the cause of epidemics fatal to cattle, 
it would not be possible to make any definite statement on the 



subject until observations can be made on the spot during the time 
when the epidemic is running its course, so that animals can be 
examined systematically, the clinical course of the disease watched 
and material obtained immediately after death. We cannot 
emphasise too much the importance of properly directed and 
sustained work in the investigation of problems of such economic 
importance; it is only by this means that the true nature of these 
ever-recurring epidemics will be elucidated. 


TRYPANOSOMIASIS 

During our visit to Sierra Leone we had the opportunity of 
examining a considerable number of domestic stock, both in the 
Colony and in the Protectorate. In all, 143 cattle were examined, 
and trypanosomes were found in nineteen. The proportion of 
infected cattle must, without doubt, be very much greater than this, 
as in the majority of cases an examination of a single fresh 
preparation of the blood was all that could be undertaken. These 
cattle, which were brought to Freetown for slaughter, appeared to 
be the most heavily infected; trypanosomes were discovered in the 
blood of fourteen out of thirty-four animals immediately after they 
had been killed at the Military slaughter-house. A probable 
Explanation of this is that so soon as animals begin to lose condition 
they are taken by the natives to be sold in the Freetown market. 
In the nineteen infected animals Z. congolense was found eleven 
times, Z. vivax twice, a double infection of T. congolense and 
Z. vivax five times, and T. gambiense once. 

The blood of ten dogs was examined, but in no cases were 
trypanosomes found. One of the animals was, however, proved to 
be infected with Z. congolense by inoculation of rats. No instance 
of infection with trypanosomes was found in the seven goats and 
seven sheep examined. 

In connection with the information derived from examination of 
domestic stock it is of interest to refer to observations on the amount 
of trypanosome infection obtaining in G. pal falls. 

Four hundred wild Glossina palpalts caught on the Cape 
Lighthouse Peninsula were dissected and examined; trypanosomes 
were found in twenty-one of them. 



4i 6 


Tabli I.—Showing the manner in which trypanosomes were recognised in infected domestic 

stock 


No. 

Animal 

Trypanosomes 
found in 
peripheral 
blood 

Animals 

inoculated 

Result of 
inoculation 

Diagnosis 

i 

Ox 11 

T. congolense 

... 

... 

7 . congolense 

z 

Ox 23 

T. vivax 

Ox I 

Ox II 

Rat 15a 

Rat 15b 

Rat 15c 

Negative 

Negative 

Negative 

Negative 

Negative 

T. vivax 

3 

Ox 88 

7 . congolense 

Rat 21 

7 . congolense 

T. congolense 

4 

Ox 89 

T. vivax 

Ox III 

Ox IV 

Rat 16a 

Rat 16b 

Rat 16c 

7 . vivax 

7 . vivax 

Negative 

Negative 

Negative 

7 . vivax 

5 

Ox 100 

Negative 

Rat 24 

T. gambiense 

T. gambiense 

6 

Ox 108 

T. congolense 

... 

... 

T. congolense 

7 

Ox in 

T. congolense 



7 . congolense 

8 

Ox 112 ... 

T. congolense 



T. congolense 

9 

Ox 120 

T. congolense 



T. congolense 

IO 

Ox 121 ... 

T. congolense 


... 

T. congolense 

n 

Ox 137 ... 

(T. congolense 
|T. vivax 

Rat 11a 

Rat nb 

Negative 

Negative 

J T. congolense 
\ T. Ct« 7 A* 

12 

Ox 140 

7 . congolense 

Rat 12a 

Rat 12b 

7 . congolense 

T. congolense 

T. congolense 

13 

Ox 141 ... 

7 . congolense 

Rat 13a 

Negative 

7 . congolense 

14 

Ox 142 ... 

7 . congolense 

Rat 13b 

Negative 

7 . congolense 

15 

Ox 143 ... 

7 . congolense 

... 

... 

7 . congolense 

16 

Ox 144 ... 

( 7 . congolense 
(T. vivax 



j 7 . congolense 
( 7 . vivax 

17 

Ox 145 ... 

| T. congolense 
( 7 . vivax 

Mi 

... 

( 7 . congolense 
( 7 . two* 

18 

Ox 146 

f T. congolense 
\T.vivax 



f 7 . congolense 
\ 7 . vivax 

19 

Ox 147 ... 

( T. congolense 
{ 7 . vivax 


... 

f 7 . congolense 
( 7 . vivax 

20 

Dog 155 ... 

Negative 

Rat 30a 

Rat 30b 

7 . congolense 

T. congolense 

7 . congolense 







4 ! 7 


Table 2. —Giving results of dissection of wild Glossina palpalis found to be infected with 

trypanosomes 


No. of fly 

Salivary 

Proboscis 

Intestine 

Result of inoculation 

glands 




« 

0 

0 

4 - 4 - 4 - 

2 rats, negative 

2 

0 

+ 4 - 

0 


3 

0 

4 - 4 - 

4 - 4 - 

1 rat, gut and proboscis contents, 





negative 

4 

0 

4 - 4 - 4 - 

0 


5 

0 

4 - 4 - 

0 


6 

0 

4 - 4 - 

0 


7 

0 

4 - 4 - 4 - 

0 


8 

0 

4 - 

0 

(1 rat, gut contents, negative 

9 

0 

4 - 4 - 4 - 

4 - 4 - 4 - 

J 1 rat, proboscis contents, died on 6th 





l 

10 

0 

4 - 

0 


11 

0 

4 - 4 - 4 - 

0 


12 

0 

4 - 4 - 4 - 

0 


13 

0 

4 * ! 

0 


14 

0 

4 - 4 - 1 

0 


>5 

0 

0 

4 - 

(i rat, gut contents, died on 6th day 

16 

0 

4 - 4 - 4 - 

4 - 4 - 4 - 

j 1 rat, proboscis contents, died on 6th 
[ day 

>7 

0 

4 - 

0 

18 

0 

4 - 

0 


>9 

0 

4 - 

0 


20 

0 

4 - + 

4 - 4 - 4 - 

(i rat, gut contents, negative 
{1 rat, proboscis contents, negative 

21 

0 

4 - 

0 



4 * Signifies trypanosomes scanty. 

4- 4 - Signifies trypanosomes numerous. 
4 - 4 * 4 * Signifies trypanosomes swarming. 








It will be seen from Table 2 that of the twenty-one flies found 
to contain trypanosomes, the proboscis alone was involved in 
fifteen, the gut alone in two, whilst there was a heavy infection of 
both gut and proboscis in four. No instance of invasion of the 
salivary glands was encountered. Assuming that these parasites 
were trypanosomes pathogenic to man and domestic stock, the result 
of previous work allows us to conclude that fifteen of the above flies 
were infected with T . vivax , four with T . congolense , and the 
remaining two with a trypanosome belonging either to the 
Congolense or the Gambiense group; which, we are unable to 
decide owing to the fact that the cycle of development of the 
trypanosome in the fly was incomplete. Inoculations into rats were 
made, with negative results in each case, from five of the flies 
found to be infected. Unfortunately in two instances (Flies 9 
and 16) the rats which had received the contents of the proboscis 
died six days after inpculation, but in two other cases (Flies 3 
and 20) the rats lived long enough to have become infected. The 
fact that these animals did not become infected does not prove that 
the trypanosome found in the flies was not T. congolense , as it has 
now been established that it is always easy to infect rats with this 
trypanosome. A similar failure of T. congolense to establish itself 
in these animals is seen in the case of three rats inoculated 
respectively from Oxen 137, 141 and 142 (see Table I), which were 
found on direct examination of the blood to be suffering from an 
infection with T. congolense . Previously the trypanosome in 
question would have been designated T . nanum y but recent 
work shows that the difference between the latter parasite and 
T. congolense is only an apparent one. 

A small number (95) of wild G. palpalis was fed on rats; no 
infection resulted. 


BABESIASIS AND THEILERIASIS 

Whilst examining blood films of cattle, species belonging to the 
genera Babesia and Theileria were seen. In about 5 per cent, of 
the animals examined a few large double pear-shaped bodies and 
an occasional large amoeboid form were encountered in the red cells. 
This parasite was undoubtedly Piroplasma bigemtnum. 



+i 9 


The other parasite occurred much more frequently, being found 
in between 20 and 30 per cent, of the animals examined. Small rod- 
and ring-shaped bodies showing distinct cytoplasm and chromatin 
were seen in the red cells. In some animals 1 or 2 per cent, of 
the cells were invaded, whilst in others only an occasional parasite 
was found. These bodies seem to be identical with those discovered 
by Macfie (1914) in cattle in Nigeria. They did not appear to 
produce any symptoms of disease. Most of the animals in which 
they were found were in perfect health; a few were emaciated, but 
in these trypanosomes were also found. 

Owing to the fact that so large a proportion of cattle were 
discovered to harbour these bodies in their blood—usually they 
were present in very small numbers only—we did not consider that 
inoculation experiments into local cattle would afford any evidence 
as to the inoculability or otherwise of the parasite. No 1 blue bodies * 
of Koch were found in the blood, or in smears made from the 
kidneys, liver, spleen, lungs, or bone marrow of six infected oxen. 
Theiler (1907) describes Theileria parva and Theileria mutans as 
being very similar in appearance. He differentiates between them 
on the following grounds. Th. parva is very virulent, is non- 
inoculable, and is characterised by the presence of the ‘ blue bodies 1 
of Koch; whereas Th. mutans is non-virulent, is inoculable, and the 
‘blue bodies' of Koch are not found. There is little reason to 
doubt, therefore, that the parasite in question is Theileria mutans. 

All the ticks which could be found were collected from each of 
the ninety cattle examined in the Warri at Batkanu; the animals 
were thrown and carefully searched. The only species encountered 
were Boophilus australis (126 adults, and 395 nymphs) and 
Amblyomma variegatum (4 adults, and 28 nymphs). We are 
indebted to Professor G. H. F. Nuttall for kindly determining 
them. 


CONCLUSIONS 

I. Cattle are not bred to any extent in the Colony of Sierra 
Leone or in most parts of the Protectorate. Bullocks are, however, 
imported from French Senegal in considerable numbers, and 
gradually find their way down to Freetown for slaughter. 


420 


2. Serious outbreaks of disease in the form of epidemics occur 
amongst the cattle of Sierra Leone. 

3. These epidemics have been attributed to various causes, but 
their real nature is still obscure. 

4. Trypanosomiasis of cattle is common. Of the thirty-four 
animals examined at the slaughter-house in Freetown, fourteen 
(41 per cent.) were found to be infected. As only a single blood film 
was examined, the real percentage of infections is certainly much 
higher. 

5. T. congolense and T. vivax are the parasites most commonly 
found. T. gambiense was met with once. 

6. About 5 per cent, of the animals examined were found to be 
infected with Piroplasma bigeminutn. 

7. Theileria mutans was encountered in between 20 and 30 per 
cent, of the cattle examined. 


REFERENCES 

Macfie, J. W. Scott (1914). Notes on some Blood Parasites collected in Nigeria. Annals 
Trop . Med . and Parasitol ., VIII, pp. 439-468. 

Theilek, A. (1905-6). Report of the Government Veterinary Bacteriologist. Transvaal 
Department of Agriculture, Pretoria. 

- (1906-7). Report of the Government Veterinary Bacteriologist. Transvaal Depart¬ 
ment of Agriculture, Pretoria. 



421 


THE ETIOLOGY OF JUXTA-ARTICULAR 
SUBCUTANEOUS NODULES 


BY 

J. B. DAVEY, M.B. Lond., D.T.M. Liverpool 
(Received for publication 16 June, 1915) 

Whilst engaged, during the months of June, July and August, 
1913, in examining the natives of the low-lying part of the Dedza 
district of Nyasaland for sleeping sickness, two facts soon forced 
themselves upon my attention. One of these was the prevalence of 
yaws and the other the frequent occurrence amongst these people 
of firm subcutaneous nodules, varying in size from that of a pea to 
that of a duck’s egg, in the vicinity of joints. Observations were, 
therefore, made on a series of 2,378 adults (964 males and 1,414 
females) and 567 children, natives of this district, and the results are 
presented in this paper, as they seem to be of interest in elucidating 
the etiology of such nodules. 

The nodules which, as I have said, vary considerably in size, 
are firm, painless, not tender, show no tendency to break down, 
and are always situated in close relationship to the subcutaneous 
portions of bone. They are, however, freely movable on the bones 
and the skin over them, except in a few cases where definite signs 
of trauma accounted for adhesion to the skin. Their favourite site 
is the posterior border of the ulna, about two inches from the tip of 
the olecranon, and they were found, in this situation, on one or 
both sides, in 55 of the 2,378 adults examined. Other fairly 
common sites are the great trochanter of the femur and the lower 
part of the patella: less commonly they were found over the 
malleoli, and in one case on the zygoma. Frequently several were 
found in one individual at these various positions. At the patellar 
site they were usually accompanied by enlarged bursa. Altogether 
in 80, out of the 2,378 adults examined, they were found in one or 
more positions. They were never found in children, of whom 567 
were examined for them. 



4 22 


These observations led me to the conclusion that the nodules are 
those referred to, in works on Tropical Medicine, as ‘ Juxta-articular 
Nodules,* and they have not, so far as I am aware, been reported 
from this part of Africa before. 

Of the 964 men, 14 were actually suffering from, and 160 showed 
clear indications of having had, yaws: of the 1,414 women, 29 and 
234 respectively: for the 567 children, the figures were 15 and 70. 

With eight exceptions, the 80 subjects of these subcutaneous 
nodules had characteristic scars of, and admitted having had, 
yaws: of the eight exceptions, seven had the typical pigmented, 
finely wrinkled scars, but denied infection. Yaws is so commonly 



Subcutaneous Juxta-articular Nodule on the forearm of a man who had yaws as a child. 


contracted by these people in infancy and early childhood that many 
may forget all about it by the time they reach adult years. The 
opinion which I hold that these subcutaneous nodules are a late 
manifestation of yaws, comparable to, but without the tendency to 
softening of syphilitic gummata, is not, I think, invalidated by the 
above-mentioned exceptions. In most cases the nodules appear 
years after the manifest lesions of yaws have healed, but in 
four cases they were found in persons still showing unhealed 
granulomata. 

For comparison, I was able subsequently to examine 327 adults 


and 266 children in another (Mlanje) district of the Protectorate 
where yaws is not, so far as I am aware, known. The numbers are, 
unfortunately, small. No recent case was found amongst these 
people, who belong to quite a different tribe from the inhabitants of 
Dedza district, and many of whom have, within recent years, 
immigrated from Portuguese territory. Only two of the adults 
admitted having had yaws or showed indications of infection: both 
of these had characteristic yaws scars and also subcutaneous 
nodules: one had also beautifully pitted hands and feet—a 
condition to which Castellani (1910) has drawn attention. These 
two persons had contracted the disease in another part of the 
country. None of the 266 children examined showed any indica- 
• tions of the disease. 

In the Journal of the London School of Tropical Medicine 
(1913), in a review of a paper by Ouzilleau, it is stated that in 
Mbomau adult Filaria volvulus were found ‘ in cysts or tumours, 
generally superficial, situated under the skin,* and ‘ the conclusion 
must be drawn that the volvulus cysts situated near joints do not 
differ markedly from those tumours known as “ Juxta-articular 
Nodules .” 9 ‘ Brumpt had observed many cases of these nodosities 

in Uganda and some on the Ouelle * ... ‘ in every case of this 
sort seen by the author he had punctured the tumour and found the 
embryos and sometimes fragments of the adult of Filaria volvulus 9 
I therefore punctured one nodule, and excised two from the forearm 
in another case: neither in the small quantity of material removed 
by puncture, nor in scrapings from the cut surface of the excised 
nodules was anything resembling a filaria seen. Microscopically, 
fragments of fibrous connective tissue but no parasites or organisms 
of any sort were found. On section the nodules were solid, tough 
and lay embedded in subcutaneous fat. 

Under the tertiary stage of yaws, Castellani and Chalmers 
mention the occurrence of gummatous-like nodules which soften and 
break down: such a condition I have observed in a few cases of 
yaws: the nodules to which I refer have no such tendency to 
softening. 

The observations recorded above point strongly, in my opinion, 
to these nodules being a late manifestation of yaws, and they so 
closely resemble the 1 Juxta-articular Nodules * described in 



Castellani and Chalmers' Manual of Tropical Medicine that there 
can be little doubt of their identity with the latter. 

Since these observations were made, I have read in the Tropical 
Diseases Bulletin (1913) that Mouchet and Dubois have observed 
similar nodules, and state that the natives in the Congo consider 
them a late manifestation of yaws. The sectional editor adds that 
Jeanselme's Juxta-articular Nodules are due to a fungus— 
Nocardia carougeaui , Brumpt, 1910—but that in Africa, nodules 
induced by Filaria volvulus may closely simulate them. 


REFERENCES 

Castellani and Chalmers (1910). Manual of Tropical Medicine , lit edition, p. 1137. 
-(1910). Idem p. 868. 

Journal of London School of Tropical Medicine (1913), Vol. II, part 2, p. 133. 
Tropical Diseases Bulletin (1913), Vol. I, No. 9, p. 530. 



425 


ANKYLOSTOMIASIS IN DOGS IN 
SIERRA LEONE 


\Being the Fifth Report of the Thirty-second. Expedition of the 
Liverpool School of Tropical Medicine , 1914-1915.] 

BY 

WARRINGTON YORKE 

AND 

B. BLACKLOCK 

(Received for publication 9 July , 1915) 


Ankylostome infection appears to be universal in dogs in 
Freetown; of seven examined by us, all were found to be heavily 
infected. More detailed examination showed that the infection was 
due to two parasites— Ankylostoma caninum , Ercolani, 1859, and 
Ankylostoma ceylanicum, Looss, 1911—which were present in the 
intestines in about equal numbers. These' species are readily 
distinguished one from the other by the characteristic arrangement 
of their teeth. The mouth of Ankylostoma caninum is armed with 
three pairs of prominent ventral teeth, whereas in Ankylostoma 
ceylanicum there is one pair of large ventral teeth, and one very 
small pair near the base of the former, but on a slightly deeper 
plane. These characters are illustrated in Figs. 1 and 2. 



Fig. 1. Mouth of Ankylostoma caninum. 
X 300. 


Fig. 2. Mouth of Ankylostoma ceylanicum. 
X 300. 



426 


The bursa of the males is very similar in the two species. It 
consists of two large lateral and a small dorsal lobe. The arrange¬ 
ment of the rays is as follows: In each lateral lobe there is an 
anterior ray which is cleft, an antero-extemal ray, a median ray 
which is doubled, and a postero-extemal ray which arises from a 
common trunk with the single posterior ray. In the dorsal lobe is the 
posterior ray, which exhibits slight differences in the two species. 
In both it is bifurcated in its terminal third, and each of the branches 
is at its extremity tridigitate. It is in the character of these terminal 
digitations that the slight difference is found (see figs. 3 and 4). In 




Fio. 3. Posterior ray of bursa of Ankylostoma Fig. 4. Posterior ray of bursa of Ankylostoma 

caninum . x 500. ceylanicum. x 300. 

both species the two inner digitations are small, being separated by 
a mere notch. In Ankylostoma caninum , the cleft sparating the 
two inner from the outer digits is shallow, but in Ankylostoma 
ceylanicum , the cleft is deep, being about half the length of the 
branch of the posterior ray. 

The average length of Ankylostoma caninum is greater than that 
of Ankylostoma ceylanicum. A few were measured by us with the 
following results: 

Ankylostoma caninum : Males 6*5-8 mm., females 7*5-14*5 mm. 
Ankylostoma ceylanicum : Males 6 mm., females 7-10 mm. 

The discovery of Ankylostoma ceylanicum in the dog in Sierra 
Leone is of interest. This species was first described by Looss in 
1911, from a civet cat from Colombo. It was subsequently found 
by Lane (1913) in the dog and cat in Bengal, and also in a lion from 
the Calcutta Zoological Garden. In the same paper Lane described 
this parasite as occurring occasionally in human beings. 

Gomes de Faria, in Brazil (1910), found dogs and cats infected 
with Ankylostoma caninum and a parasite which he described as a 




4 2 7 

new species under the name Ankylostoma braziliense 9 but Leiper 
(1913)* from a comparison of figures published by Lane and 
de Faria, concludes that Ankylostoma braziliense and Ankylostoma 
ceylanicum are identical. 

We had no opportunity of determining whether Ankylostoma 
ceylanicum occurs in human beings in Sierra Leone, where infection 
with Ankylostominae is exceedingly common. The importance of 
ascertaining whether this species occurs in man is obvious. If 
Ankylostoma ceylanicum is found in human beings in Freetown, the 
dog reservoir of the infection is a factor which must be borne in 
mind when prophylactic measures are under consideration. 


CONCLUSIONS 

Dogs in Freetown are heavily infected with Ankylostominae. 
The species found were Ankylostoma caninum and Ankylostoma 
ceylanicum; they were present in about equal numbers. 


REFERENCES 

de Faria, G. (1910). Contribution towards the classification of Brazilian Entozoa. Memorial do 
Instituto O steal do Cruz , II, pp. 286-293. 

Lank, Clayton (1913). Tricbostrongylus columbriformis (Giles, 1892), a Human Parasite. Ind. 
Med. Gaz ., XLVIII, pp. 129-132. 

-(1913). Agcbylostoma ceylanicum , a New Human Parasite. Ind. Med. Gaz ., XLVIII, 

pp. 217-218. 

Lkipxr, R. T. (1913). The Apparent Identity of Agcbylostoma ceylanicum (Looss, 1911) and 
Agcbylostoma braziliense (Faria, 1910). Journ. Trop. Med. & //yg., XVI, pp. 334-335. 

Looss, A. (1911). Records of the School of Medicine , Cairo, Vol. IV. 




4 2 9 


STUDIES IN BLACKWATER FEVER* 

IV.—NOTE ON A CASE OF QUARTAN 
MALARIA ASSOCIATED WITH 
BLACKWATER FEVER 

BY 

J. W. W. STEPHENS 

SIR ALFRED JONES PROFESSOR OF TROPICAL MEDICINE, THE UNIVERSITY OF LIVERPOOL 


(Received for publication 2 July , 1915 ) 

One Chart 

Past History 

X.Y.Z. European, W. Africa. Length of residence in Africa 
four tours, t 

31.3.13. —6.4.13. Sunstroke. 

29.4.13. — 3 . 5 . 13 . Sciatica and malaria. Has been ill with 
more or less fever for last seven days 

Present History 

12.8.13. 4 p.m. Admitted to hospital. Condition : Skin yellow, 
unhealthy looking; conjunctivae pale; gums pale; tongue dry 
yellow-brown fur. Liver palpable i^* below costa. Pulse, 108, 
Temperature, ior8°. 

14.8.13. Patient progressing favourably until 

15.8.13. at 4 p.m., feeling of distress, anorexia, frequent shivers, 
inclination to vomit. 10.30 p.m. Blackwater passed. 

16-19.8.13. Frequent vomiting, extreme jaundice, increasing 
heart-failure. 

18.8.13. Slight oozing of blood from gums. 

19.8.13. Profound collapse. Death 1.20 p.m. 

• Part I j Annals of Trop. Med. & Parasitol 1913. Part II: Ibid., 1914. 

Part III : Ibid., 1915, p. 201. 


f A tour is a year. 



43 ° 

In the accompanying tables I have given an abstract of the 
available data as to the case. They are presented here for the sake 
of completeness rather than for any bearing they have on the 
problem raised. 


Record of Quinine 

12.8.13., 4 P* m * Quinine grs. vi. intramuscularly. 

13.8.13. ‘Mist, quin.' quartis horis (at what hours given not 
stated). 

14.8.13. Presumably ‘Mist, quin.' continued. 

15.8.13. Presumably continued during the day, ? till 4 p.m.. 
as it is stated that in the evening it was discontinued, but whether 
before or after the blackwater is not stated. Aspirin 10 grs. 
presumably at or after 4 p.m. 

(NOTE. —The Mist, quininae of the B.P.C. contains gr. i. to 31, 
but whether the above mixture is this, it is impossible to say.) 


Blood 



Red cells 

Hgb. 

Total 

count 

leucocytes 

Percent¬ 

age 

count 

number 

counted 

Urge 

mono¬ 

nuclear 

Lympho¬ 

cytes 

Poly¬ 

nuclear 

Eosino¬ 

phil 

Parasites* 

12.8.13 ••• 









Quartan parasites, 

13.8.13 ... 

... 

... 

... 

... 

... 


... 

... 

scanty rings 

14.8.13 ... 

... 

... 

••• 

... 

... 





15.8.13 ... 

m.) 

2,500,000 

... 


300 

6 

it 

83 


Parasites negative 

(10.30 p. 
16.8.13 ...; 

5 °% 

(Tallqvist) 

... 

... 

... 

... 

1 


17.8.13 ... 

1,400,000 

3 °% 

- 

... 




' 

... 


18.8.13 ... 

6oo,ooof 

2 5 %t 

18,000 

300 

H 

15 

65 

6 

Parasites negative 


* Five Blood examinations were made, but the dates of two of them are not stated, 
t Non.—These data imply a color index of 2*i 1 




43i 


UrIKE 


Date 

Time 

Quantity 

Reaction 

Colour 

Hgb. 

Casts, etc. 

. 

... 

... 


... 

No alb. 

Bile present 

14.S.13 . 

1 5 - 8 -'3 . 

Apparently 

... 1 ... 1 

no urine passed before 10.30 p.m. 



5 ) . 

10.30 p.m. 

220 c.c. 

Faintly alkaline 

Black 

Met. Hgb. 

Granular casts 

16.8.13 . 

2 a.m. 

70 c.c. 

Faintly add 


»» 

ss 

» . 

3 a.m. 

14 c.c. 

Add 

Rather lighter 


yy 

IS . 

9 a.m. 

145 c.c. 

Aik. 

... 

ss 

ss 

n . 

9.40 a.m. 

170 c.c. 

(In the motion) 

... 



>s . 

12.15 p.m. 

240 c.c. 

Aik. 


Met. Hgb. 

yy 

» . 

6 p.m. 

240 c.c. 

ss 

Much lighter 

yy 

yy 

17.8.13 . 

12.30 p.m. 

278 c.c. 

Add 

Darker 

yy 

No casts 

» . 

— 

230 c.c. 

Aik. 

» 

yy 

yy 

18.8.13 . 

2 a.m. 

360 c.c. 

Acid 

Clearing 

yy 

yy 

ss . 

11.45 a.m. 

50 c.c. 

ss 

Amber 

yy 

■ 

yy 

ss . 

10.30 p.m. 

155 c.c. 

ss 

Light amber 

No alb. 

Trace of bile 

» . 

11 p.m. 

20 C.C. 

ss 

n 

yy 

yy 

1 

19.8.13 . 

12.45 a * m * 

60 c.c. 

yy 

yy 

yy 

, 

yy 

ss . 

12 noon 

? 

ss 

... 

... 

No casts 


Post-mortem (partial) Fite Hours apter Death 


Skin . 

Very yellow 


Liver . 

Very large, yellow mottled, 

Fatty degeneration, perilobular 


soft, old adhesions 

fibrosis. No parasites, some 
pigment 

Spleen . 

Large 

No parasites, some pigment 

Stomach. 

Petechiae oozing blood, con¬ 
tents green bile 


Small intestine. 

Petechiae oozing Mood 


Gall bladder . 

Yellow bile 

j 

Kidneys . 

Large, pale 

Many lobules blocked with 
‘ haemoglobin ’ casts 

Bone marrow . 

Pale 

No parasites, some pigment 























+ 3 * 


This case is of interest for the following reasons. 

1. Immediately after the blackwater, 15.8.13, 10.30 p.m., no 
malaria parasites or pigment were found, i.e., there is no evidence 
of a malaria infection, yet on 12.8.13, 4 P-m., scanty quartan 
parasites were found, i.e., the patient at that time was infected with 
malaria. The records do not permit of any conclusion as to when 
evidence of malaria infection ceased, for although two other blood 
examinations (presumably negative) were made, yet it is not stated 
when they were made, i.e., whether before or after the blackwater. 
Further, the only evidence of a malaria infection post-mortem is the 
presence of pigment, parasites being absent. It is improbable, 
however, that a quartan infection was got rid of by three days’ 
quinine treatment. We have then a case of blackwater fever, 
negative as regards malaria, subsequent to the blackwater, but 
positive three days previously, and also positive (pigment) post¬ 
mortem. 

2. The diagnosis of quartan parasites on the 12th is supported 
by the character of the chart (see p. 433), which is quartan in 
character. If we assume that the rise on the 15th is a quartan 
paroxysm, then we have the actual paroxysm in this case, for some 
reason or other, associated with blackwater. It may be argued, 
however, that the paroxysm on the 15th is due to the onset of black¬ 
water per se y and that it is a coincidence that it has occurred when 
we should have naturally expected a quartan paroxysm. This 
possibility cannot be disproved, but also the possibility—if nothing 
more—exists that this is not so, and that in this case a quartan 
paroxysm is closely concerned with the blackwater attack. The 
chances appear to me to favour this interpretation, because the 
quartan paroxysm would occur on this day, whereas the blackwater 
might have occurred at any other time. A study of temperature 
charts (unfortunately few are available), previous to the onset of 
blackwater, would probably throw light on this point. 

I hope, in due time, to be able to collect a number of suitable 
cases. I have to acknowledge with pleasure the kindness of the 
Colonial Office in putting the records of this case at my disposal. 










































Volume IX 


December, 1915 


No. 4 


ANNALS 

OF 

TROPICAL MEDICINE AND 
PARASITOLOGY 


ISSUED BY 

THE LIVERPOOL SCHOOL OF TROPICAL MEDICINE 


Edited by 

Professor J. W. W. STEPHENS, M.D. Cantab., D.P.H. 

Professor R. NEWSTEAD, M.Sc., J.P., F.R.S., A.L.S., F.E.S., Hon. F.R.H.S. 
Professor WARRINGTON YORKE, M.D. 

AND 

Professor Sir RONALD ROSS, K.C.B., F.R.S., M.D., F.R.C.S., 

Major I.M.S. (Ret.) 


Editorial Secretary 
Dr. H. B. FANTHAM, 

School of Tropical Medicine , 
The University , 
Liverpool. 



C. 'Tinling & Co. t Ltd. 

Printers to the University Press of Liverpool 
$3 Victoria Street 



435 


NUCLEAR VARIATIONS OF THE 
NEUTROPHILE LEUCOCYTES 
(ARNETH COUNTS) IN MALARIA 
AND YELLOW FEVER 


BY 

J. W. SCOTT MACFIE, D.Sc., M.B. 

WEST AFRICAN MEDICAL STAFF, GOLD COAST 

(Received for publication 22 July , 1915) 


Considerable interest has recently been evinced in the changes 
observed to take place in various diseases in the nucleus of the 
polymorphonuclear leucocyte; and, following Ameth’s method of 
classification of these cells, it has been shown that there is an increase 
of the cells of Classes I and II in many microbic diseases. The 
researches of Chamberlain in 1910 first drew attention to the fact 
that a similar phenomenon is exhibited by persons living in the 
tropics; and it has recently been suggested by Breinl and Priestley 
(1914), as the result of their examinations of school children in 
Queensland, that this change may be ‘ due to the effects of a tropical 
climate upon the white race living in the tropics/ 

An increase of the percentage of polymorphonuclear leucocytes 
belonging to Classes I and II has been observed in a great many 
different diseases. It occurs, according to Cooke (1915), in 
typhoid, scarlet fever, diphtheria, measles, chicken-pox, erysipelas, 
tonsillitis, pneumonia, whooping-cough, puerperal fever, tuber¬ 
culosis, nasal catarrh, and septic conditions, and probably in other 
diseases also. It is important, therefore, in considering the 
significance of a shift to the left of the Arneth count, such as that 
found in Europeans in the tropics, to determine in the first instance 
the effect on the polymorphonuclear leucocytes of the diseases 
prevalent in these climates, and especially that of such an infection 



43 ^ 


as malaria which may lie latent in the body for long periods without 
exciting active symptoms. In the following pages are embodied the 
results of the examinations of a number of cases of malarial fever 
and yellow fever with special reference to the changes occurring in 
the polymorphonuclear leucocytes. 

The blood-films were all stained by Leishman's method. Two 
hundred successive neutrophile leucocytes were examined in each 
film, each hundred being counted separately and the two sets of 
figures being accepted only if they agreed approximately. Nuclear 
fragments connected by a fine thread were counted as separate; but 
those joined by a definite band were counted as one. In all 
doubtful cases the cells were referred to the higher of the two classes 
to which they might have been assigned. 

In making Arneth counts there must always be a very consider¬ 
able personal variation. Many cells are exceedingly difficult to 
assign to any particular class, and in such cases different investigators 
might easily differ in their ultimate decisions. The various 
estimates of the normal Arneth index, which range from twenty-five 
to fifty-six, are in themselves a proof of this fact. Allowance must 
always be made, therefore, for some degree of individual difference 
in comparing counts made by different observers, and too great 
importance should not be attached to small degrees of shift. A 
series of counts by the same observer in different conditions should, 
however, be strictly comparable. In making the counts referred to 
below, great pains were taken to try and follow the course of the 
nucleus before attempting to decide to which class the cell should 
be assigned. It was assumed that the nucleus was a continuous 
body with two free ends, and that it should be possible to trace the 
connexions between the different lobes. This assumption was not 
always borne out in fact, as leucocytes are occasionally met with 
even in normal blood in which the nuclear fragments are actually 
separate from one another. Such cells are probably dead cells; but 
as they are invariably easy to classify, they do not affect the counts. 
In my experience the greatest difficulties occurred in films which 
showed a normal, or nearly normal, count. In those in which there 
was a marked shift to the left, and these were the most interesting 
of the series, there was seldom any doubt whatever as to the classes 
in which the various leucocytes should be placed. 



I. THE RESULTS OF ARNETH COUNTS IN HEALTHY 
EUROPEANS AND NATIVES 


A. Europeans. 

Twenty-nine apparently healthy Europeans were examined at 
various stages of their tours of service in West Africa (see Table I). 
The average Ameth index was 51 *6; that is, there was a slight shift 
to the left in the count of about the same degree as that found by 
Chamberlain and Vedder (average index 46*2) in seventy-two healthy 
American soldiers in the Philippine service. 

Eleven of the subjects had been in residence in the Gold Coast 
for less than a year, and eighteen for twelve months or longer. The 
Ameth index in the former averaged 50 6, and in the latter 516; 
a scarcely perceptible increase in favour of those who had been in 
West Africa for the longer period. 

Two of the Europeans had landed at Accra only one and four 
days respectively before they were examined. In the latter the 
Ameth count was practically normal (index 41*5), but in the former 
there was a slight but distinct shift to the left of the count for which 
there was no apparent cause. He had previously served in West 
Africa, and had, of course, been in the tropics during the last few 
days of the voyage before he landed, and had possibly, although 
not probably, been infected by mosquitos. But in this connexion 
it should be noted that it is not a very uncommon occurrence for men 
returning to West Africa to show symptoms of malaria soon after 
landing; and that this is probably due to their having failed, during 
their leave of absence, to eradicate from their systems the malarial 
infections contracted during their previous tour of service. 

In a subsequent part of this paper it is shown that in malaria 
there is a pronounced shift to the left of the Ameth count, and that 
this phenomenon is observable before the parasites are sufficiently 
numerous to be found in the peripheral blood, and that it may 
persist for a considerable time after the patient has apparently 
recovered. Practically all the apparently healthy Europeans 
examined had suffered from malarial fever at some time during their 
tour, and it may be assumed that they had all been repeatedly 
inoculated with malaria parasites by infected mosquitos. I believe 
that this is a sufficient explanation of the slight shift to the left of 




Arneth classification per cent. 


No. Sex Length of tour __Arneth 


I 

11 

III 

IV 

V 

Index 

1 

M. 

4 days 


5*5 

36-0 

41-0 

* 5*5 

2*0 

4**5 

2 

M. 

1 

day 


8-5 

45-0 

39 -o 

7*o 

o*5 

53*5 

3 

F. 

3 months... 


6-o 

3 °*° 

47*5 

14*0 

2 *5 

36-0 

4 

M. 

6 



I o-o 

29*0 

37 *o 

21*0 

3 *o 

39 *o 

5 

M. 

6 

« - 


5 -o 

41*0 

39*5 

* 3*0 

**5 

46-0 

r> 

F. 

7 



,2 *5 

38-5 

34*5 

,2 *5 

2*0 

51-0 

7 

M. 

10 

1 ? 


9*5 

4**5 

37*5 

11*0 

0-5 

51*0 

8 

M. 

10 

11 


* 3*5 

47 -o 

30*0 

9*0 

o*5 

60*5 

9 

F. 

10 

„ ... 


16-o 

47 -o 

2 9*5 

7*5 

— 

63*0 

IO 

M. 

11 

,, ... 


17-0 

38*5 

36*5 

8-o 

— 

55*5 

11 

M. 

11 

V 


* 3*5 

46-5 

36-0 

4 -o 

— 

6o*o 

12 

M. 

12 

11 


15-0 

3 8-8 

35*o 

10*0 

1*2 

53*8 

13 

M. 

12 

11 


22-0 

46*0 

29-0 

3*0 

— 

68*0 

14 

M. 

12 

11 


130 

45 *° 

37*5 

4*5 

— 

58-0 

*5 

M. 

12 

T) 


17-0 

40*0 

35*o 

8 -o 

— 

57 *o 

16 

M. 

12 

11 


7-5 

4**5 

36*0 

14-0 

1*0 

49-0 

*7 

F. 

12 

11 


4-0 

41-0 

42-0 

130 

— 

45 -o 

18 

M. 

12 

11 


i 10*0 

4 2 *5 

37*o 

9*5 

1*0 

| 5 2 ‘5 

*9 

F. 

12 

5J 


j 16*0 

j 

39*5 

33*5 

11*0 


55*5 

20 

M. 

13 

?) 


18-0 

37*o 

33*5 

IO '5 

1 *0 

55 -o 

21 

M. 

13 

n 


7*5 

1 

44.0 

40*0 

8 -o 

o*5 

5**5 

22 

M. 

13 

M 


18-0 

57 *o 

2 3’5 

*•5 

— 

75 *° 

2 3 

F. 

13 

» 


9*5 

43*5 

37*5 

9*5 : 

— 

53 -o 

2 4 

M. 

13 



135 

41*0 

36*0 

9 *o 1 

t 

0-5 

54*5 

2 5 

M. 

14 

J? 


7 *o 1 

33*o 

41*0 

i 7 *o | 

2*0 

40*0 

26 

M. 

14 

V 


8 -o 

38*5 

4 2 *5 

10*0 ' 

1*0 

46-5 

2 7 

M. 

»5 

11 


5*5 

35 *o 

43 *o 

1 

14,5 

2*0 

40*5 

28 

M. 

15 

1) 


8 -o 

2 4*5 

40*0 

i 

2 3*5 

4*0 

3 2 ’5 

2 9 

M. 

16 

11 


15-0 | 

37 *o 

35*o 

IPO 

2*0 

52*0 

Averages 

| 

u 4 ; 

1 

40*2 

367 

107 

1*0 

51-6 












439 


the Ameth count found in these subjects. I do not consider that 
there is at present sufficient evidence in support of the view that this 
phenomenon is due to the influence of a tropical climate on the white 
race. The fact that the shift to the left was almost the same in those 
that had been in West Africa over a year, and in those who had but 
recently arrived, is, I think, contrary to any such supposition. 

It is clear, at any rate, from these examinations that certain 
individuals may show a normal Arneth count after more than a year’s 
residence in West Africa (see Table I, cases Nos. 25 and 26). Some 
of the individuals in whom the count approximated most nearly to 
the normal were men who had spent many years on the 4 Coast,’ and 
who might be supposed to have become acclimatised, or to have 
mastered the conditions of life most suitable for West Africa. Others 
were individuals who were on the whole better cared for than the 
majority of their fellows, who had better quarters, and who were to 
a great extent relieved from the petty domestic worries that are 
responsible for so much irritation, and probably so many of the 
ailments, in West Africa. 

B. Natives. 

Ameth counts were made in the cases of twenty apparently 
healthy natives at Accra. All the blood films were taken on the 
same day, and within the same hour, from twenty labourers who 
were actually at work at the time when they were called up for 
examination. The average index worked out at 55*92, and the 
averages of the percentages of the polymorphonuclear leucocytes 
belonging to the five classes of Arneth’s classification showed a slight 
shift to the left (see Table II). 

There were, however, great variations in the different counts; 
and the index ranged from 26'5 to 98*5. It is doubtful, therefore, 
if an average based on such a small number of counts is of any value. 
A better idea of the actual conditions found is obtained, I believe, 
by distributing the indices into groups, and plotting a curve to show 
the percentages of the cases that fall into each group. This has 
been done in Chart I. It will be observed that the crest of the 
curve in natives shows a slighter degree of shift to the left from the 
normal point than that in Europeans; although in the former the 
average index worked out at 55*92, and in the latter at 51‘6. 



440 


Table II.—Arneth counts in apparently healthy natives at Accra, West Africa. 


1 

No. , 


Arneth classification per cent. 


1 

Arneth 

index 

i 


I 

II 

III 

, IV 

V 

I 

7.0 

44*5 

00 

6 

, 9*5 1 

1*0 

; 51*5 

2 

10*0 

42-0 

41*0 

7-5 i 

°*5 

! 5 1- ° 

3 

24*0 

52*0 

22*0 

2*0 j 

— 

1 76*0 

4 

22*5 

51-0 

24*0 

2,5 ! 

— 

73*5 

5 

95 

37*5 

39*5 

12-5 ! 

1*0 

47 -o 

6 

] ' 5 '° 

36-0 

39*5 

9’5 

— 

51-0 

7 

2 ‘5 

24*0 

! 48*0 

2 1 *5 

4 *° 

26-5 

8 

10*0 

37 *° 

1 

4 °'° 

* 3 ° 

— 

47 -o 

9 

13*0 

37 '° 

1 37'5 

12-5 

— 

50*0 

10 

8 0*5 

i8*o 

I'O 

— 

°’ 5 # 

98-5 

11 

12-5 

36*5 

00 

6 

1 I'O 

2-0 

49.0 

12 

10*0 

36-5 

38-5 

130 

2-0 

46'5 

n 

8-o 

34 ’° 

l 

40*0 

17*o 

1*0 

42*0 

14 

16*o 

4 I *5 

33*5 

8-o 

1*0 

57*5 

*5 

* 4-5 

49-0 

i 3°*5 

6 *o 

1 - 

63-5 

16 

i 8‘5 

48*0 

29-0 

4*5 

1 

66-5 

17 

! 4'5 

35*5 

00 

Wl 

I 1*0 

1 

°*5 

i 50*0 

18 

i o *5 

35*0 

43-5 

105 

°'5 

1 45*5 

! 

19 

! 3-5 

36-5 

39-5 

9*5 

1*0 

i s°*° 

20 

3**5 

43-5 

23-0 

2*0 

— 

j 75 *° 

17-17 

3*75 

1 34 * 22 

9-15 

075 

! 55 ' 9 Z 


• This leucocyte was probably a dead one undergoing chromolysis. I am unable to 
account for the very high index in this case. The man when seen again a day or two later 
appeared to be in good health, and no information could be elicited excepting that he had 
had 4 fever a short time ago/ His Arneth index four days later was 64-5. 




The shift to the left of the Arneth count in natives at Accra was 
only a slight one. It was much less pronounced than that discovered 
by Chamberlain and Vedder in Filipinos (average index 65*8). 

There are several factors in the normal life of healthy natives 
that may have a bearing on the condition of the blood, especially 
the prevalence of malarial and intestinal infections. 



Chart I.—The distribution by groups of the Arneth index in (A) twenty 
healthy natives, and (B) twenty-nine healthy Europeans at Accra. 


It may be assumed that nearly every native harbours intestinal 
parasites, since ova of various species of worms, Trichomonas , and 
spirochaetes are very generally found in the faeces, and adult worms 
are recovered at practically every post-mortem examination. Of the 
blood changes produced by these infections anaemia and eosino- 
philia are the best known; but Knapp (1915) has recorded that he 








has found a shift to the right of the Arneth count in amoebic 
dysentery and ankylostomiasis. 

With regard to malaria, plasmodia are to be found in the blood 
of all, or almost all, native children; and every native in the course 
of his normal daily life must be repeatedly inoculated by infected 
mosquitos. I believe that in natives, as in Europeans, the shift to 
the left of the Arneth count found in apparently healthy individuals 
may in most cases be accounted for by such malarial infections. 
Other diseases may, of course, be responsible in some cases; such, 
for instance, as tuberculosis, or the mild attacks of yellow fever that 
scarcely incapacitate the natives at all. But as malarial infections 
dominate the field in tropical medicine to so great an extent, and 
as in this disease there is so pronounced a shift to the left of the 
Arneth count, it is not unreasonable to suppose that in the majority 
of cases the shift in apparently healthy individuals is due to this 
cause. 

If infections with intestinal parasites (amoebae, worms, 
Trichomonas , spirochaetes, &c.) are proved to cause a shift to the 
right of the Arneth count, there must be a condition of balance in the 
cases of most natives in which the shift to the left resulting from 
malarial infections on the one scale weighs against the shift to the 
right due to intestinal parasites on the other. And the same 
condition may occur in latent infections with syphilis and leprosy, 
diseases in which Knapp also noted a shift to the right. 


II. THE RESULTS OF ARNETH COUNTS IN MALARIAL FEVER 

In a previous note (1915) it was pointed out that in malarial 
fever there is a well marked shift to the left of the Arneth count, and 
that this phenomenon persists after all parasites have vanished from 
the blood, and after the patients have recovered from the attacks. 
At the time this preliminary note was published I had not seen any 
other recorded observations on the nuclear variations of the poly¬ 
morphonuclear leucocytes in malarial fever. Recently, however, 
I have received a copy of the Indian Medical Gazette (March, 1915) 
containing a paper by Knapp on ‘ The Significance of Arneth’s 
Leucocyte Count,’ in which the author records that he has found a 



JXtcv>r<\joy>VvJjL £4^coc^Vn sitJtow^cv^ h) CoCK ctaoo. 


443 



Chart II.—The average Arneth count in (N) twenty healthy natives, 
and (E) twenty-nine healthy Europeans, at Accra, compared with 
the normal count in Europe (Arneth) 










444 


decided shift to the left in malaria. Knapp states that 4 Out of 
21 cases, it was marked in 4, definite in 13, and absent in only 4/ 
that it was, on an average, 4 equal in tertian and subtertian fevers, 
and a single case of quartan gave a similar finding 1 ; and he 
concludes that 4 on the whole it may be fairly said that absence of 
the left-shift is presumptive evidence against malaria.’ Particulars 
of the counts are given in only two cases: the one, a case of 4 Double 
Tertian,’ had an index of (?) 669, and the other, 4 Subtertian,’ an 
index of 59*4. 

During the last few months I have performed Arneth counts in 
twenty-three cases of malaria in which the diagnosis was confirmed 
by the finding of the parasites, nine cases of fever in which there was 
presumptive evidence of malaria, namely an increase of the large 
mononuclear and transitional leucocytes to 15 per cent, or over, and 
twelve cases of fever suspected of being malarial but in which neither 
positive nor presumptive evidence of infection could be found. 
That is, counts have been made in forty-four cases of proved or 
suspected malarial fever. 

Of the twenty-three cases in which malaria parasites were found, 
ten were Europeans, one a Syrian, one a half-caste child, and eleven 
natives. In each case a very marked shift to the left of the Arneth 
count was found (see Table III). My results were, therefore, much 
more uniform than those of Knapp, and the shift observed was much 
more pronounced. In the Europeans the index averaged 84 25, and 
in two cases was as high as 95*5. In the natives it was even higher; 
the average being 897, and the highest 97*5. P. falciparum was the 
only species of parasite present in nineteen of the cases, and 
P. malariae in two. One case showed both P. falciparum and 
P. malariae , and one P . falciparum and P. vivax (see Table III). 

A rough estimation of the number of parasites present in the 
peripheral blood was made in each case, and the results are shown 
in the Table as a ratio of the number of red corpuscles to each 
parasite. An examination of the figures shows that there is no 
direct relationship between the number of parasites present in the 
peripheral blood and the degree of deflexion to the left of the Arneth 
count. The deflexion was greatest in those cases in which the 
constitutional disturbances were greatest; and, in the case of natives, 
in children. 



445 


Table III.—Ameth counts in twenty-three cases of malarial fever. 


No. 

Race 

: 

Arneth classification per cent. 

Nature of the infection ! 

Ameth 



I 

! 

i 

n 

III 

IV 

V 

1 index 

i 

European ... 

P. falciparum , 1:1-4 R*B.C. A few P. 78*0 
malariae. Fatal case 

I 7'5 

4-0 

o -5 

— 

95*5 

2 

11 

P. falciparum , 1 : 700 R.B.C. .51*0 

I 3 2 ’° 

16*o 

PO 

—■ 

00 

U 4 

6 

3 

11 

P. falciparum , 1 : 625 R.B.C. .22*0 

53 *o 

23-0 

2*0 

— 

75 -° 

4 

11 

P. falciparum , 1 : 250 R.B.C. .27*0 

47.0 

1 

24-0 

2*0 

— 

74 -o 

5 

11 

P. falciparum , 1 : 700 R.B.C. .27*5 

5°*5 

* 9*5 

2 '5 

— 

'vj 

00 

6 

6 

11 

P. falciparum , very rare. Blackwater fever. 6o*o 
Fatal 

27*0 

1 

12*0 

PO 

— 

87-0 

7 

11 

P. falciparum , 1 : 714 R.B.C. .56*5 

39-0 

4 -o 

°*5 

— 

95*5 

8 

11 

P. falciparum , 1 : 830 R.B.C. .43-5 

46-5 

10*0 

— 

— 

90-0 

9 

»> 

P. falciparum , few. Blackwater fever, first 29*5 
day. Mild case 

45-0 

2, ‘5 

4 -o 

— 

74*5 

IO 

11 

P. falciparum , 1 : 8500 R.B.C. . 41*0 

49*0 

90 

PO 

— 

90-0 

" 

Syrian 

P. falciparum , 1 : 1250 R.B.C. . 24*0 

56'0 

17*o 

3 -o 

— 

8o-o 

12 

Half-caste ... 

P. falciparum , 1 : 20 R.B.C . 30*0 

53-5 

16-o 

o -5 

— 

83*5 

>3 

Native 

P. falciparum, 1 : 400 R.B.C. . 67*0 

! 2 5 *° 1 

8-o 

— 

— 

92*0 


11 

P. falciparum, 1 : 713 R.B.C. . 46*0 

1 43 *° 

II-O 

— 

— 

89*0 

>5 

» 

P. malariae, very rare .j 24*0 

5 2 *5 

21*0 

2 * 5 

— 

76-5 

16 i 

11 ••• 

P. malariae , scanty . , 57*0 

34 *o | 

9 *o 

— 

— 

91*0 

17 

,, 

1 

P. falciparum (crescents only), 1 : 5000 ; and 32*0 

P. vivax, 1 : 4100 R.B.C. 

1 

55 *° 

* 3 *° 

— 1 

— 

8 7 -o 

iS j 

?? 

P. falciparum, 1 : 100 R.B.C. .35-0 

48-0 j 

16*o 

PO , 

— 

83-0 

19 

11 

P. falciparum, 1 : 19 R.B.C. 1 54*5 

35 *o 

10*0 

-5; 

— 

89-5 

20 , 

11 

P. falciparum, very few .56-0 

4 i -5 

2 ’5 

1 

— 

■ 97-5 

21 

11 ••• 

P. falciparum, scanty . 39*5 

53 *o 

7*5 

— 1 

— 

9 2 ’5 

22 

11 

P. falciparum, very rare .64*0 

33 *o , 

3 -o 

1 


i 97 *° 

23 

11 

P. falciparum, 1:91 R.B.C.54-0 

37*5 

7*5 

PO | 

— 

9 p 5 

Averages of the counts in the twenty-three cases . 1 44*3 

4 2 *3 

12-4 

PO | 

_! 

— 

86-6 







446 


The shift to the left was also well marked, but less in degree, in 
those cases of fever in which there was only presumptive evidence of 
malarial infections (see Table IV). The index averaged 79*3 in the 


Table IV.—Ameth counts in cases in which no malaria parasites could be found, but in which 
the large mononuclear and transitional leucocytes numbered 15 per cent, or over. 



Arneth classification 

per cent. 


Differential counts 

per cent. 

Race 






Arneth 

0 

-5 u 

S 

>. 

o’Sjs ! 
g S 2 i 

eft 

J V 



I 

II 

1 

III 

,v 

V 

index 

Polymorp 

nuclea 

1 

s 

J 

= S;= 
&|Sj 

1 

_c 

OU 

0 

c 

•a 

0 

w 

0 

u 

rt 

% 

European 

59 -o 

36*5 

4‘5 

— 

— 

95*5 

74 -o 

IPO 

* 5 *° 1 


— 

M 

39*5 

47-0 

I 2*0 

'•5 


8rt- 5 

6P5 

20*5 

, 6 - 5 ; 

PO 

o -5 

33 

3 i *5 

50*0 

17*0 

i *5 

— 

81-5 

5PG 

31*5 

16*5 1 

PO 

— 

51 ••• 

34 *o 

50*0 

15-0 

1*0 

— 

84-0 

4 2 "5 

37*5 

i8-o , 

p 5 

o -5 

,, ... 

8-o 

5°*5 

| 3 »*° 1 

10*0 

°*5 

58-5 

56*0 

22*5 

.8-5 

2-0 

PO 

33 ••• 

24-0 

3^*5 

l 28-5 

9’5 

p 5 

60-5 

54 -o 

l 4 -0 

! 

20*0 | 

12*0 

— 

” "1 

33 -o 

49 *° 

1 

i To 

1*0 

| 1 

1 

82*0 

66-o 

12*0 

2I *5 1 

1 

o -5 

— 

33 

33*5 

46-0 

j T 5 

3 -° 


79*5 

59*5 

6*o 

34*5 ; 

— 

1 - 

Native 

40-0 

46*0 

1 

I 3 ’° , 

PO 1 

1 

I ; 

86*o 

59 -o 

25-0 

1 5 '° 1 

o -5 

! 

! 0-5 

Averages 

33-6 

457 

1 T 3 ' 

1 1 

3-2 i 

0*2 

79*3 

58-2 

20-0 

1 9*5 

I 

2*0 

i °-3 


nine cases, and ranged from 58*5 to 95 5. No direct connexion 
between the percentage of the large mononuclear leucocytes and the 
degree of the shift could be traced. This was not to be expected, 
as the blood films were taken at different stages of the fever, and it 
is well known that the percentage of large mononuclear cells is 
subject to great fluctuations in the course of the disease. 

Of the other twelve cases of fever, seven were in Europeans, and 
five in natives. The seven cases in Europeans were all diagnosed 
as malarial from their clinical appearances, but no proof of infection 
could be obtained by the examination of the blood. In West Africa, 
where practically every European takes five grains of quinine daily, 
it is often extremely difficult to find evidences of malarial infections 
in the blood in cases of fever which look like malaria, and which 







447 


respond favourably to treatment with somewhat larger doses of 
quinine. There can be little doubt that the majority of such cases 
are actually malarial; but they are a source of difficulty and anxiety 
to the physician, and any sign that would assist in the diagnosis 
would be of great value. It is interesting, therefore, to note that in 
all these seven cases there was a decided shift to the left of the 
Ameth count. The index averaged 71 ‘35; the lowest being 61 5, 
and the highest 82 0 (see Table V). 

Table V.—Ameth counts In cases of fever suspected of being malaria, but in which neither 
positive nor presumptive evidence could be found. 

j Ameth classification per cent. 


No. 

Race 

1 

II 

III 

IV 

V 

Ameth 

index 

A 1 

European 

1 7*5 

39 -o 

32-5 

1 o-o 

1-0 

56-5 

2 

» . 

39 *o 

41-0 

18-o 

2-0 

— 

8o-o 

3 

.. 

i8-o 

48-5 

29-0 

4‘5 

— 

66*o 

4 


33 -o 

49-0 

16-o 

2-0 

— 

82-0 

5 

» . 

15-0 

46 -5 

29*5 

9 -c 

— 

61-5 

6 

” . 

23-0 

5»*5 

23-5 

2*0 

— 

74*5 

7 


30-0 

49-0 

18*0 

3-0 

— 

79.0 

Averages 

i 


: 25-07 

46-36 

2379 

4-64 

0-14 

7 * *4 

B 1 

| Native . 

i 

j 37-5 

1 

49*5 

12-0 

1-0 

— 

87-0 

2 

11 ... ... ... 

1 4*0 

28-0 

50-0 

15-0 

3 *° 

32-0 

3 

. 

16-o 

; 53 -o 

26-0 

5-0 

— 

69-0 

4 


16*5 | 

| 

54.0 

2 7 -5 

2-0 

— 

70-5 

5 

» . 

7-0 j 

44 *o , 

41-0 

8-0 

— 

51-0 

] 

Averages 


16*2 

t 

457 

3**3 

6-2 

o-6 

61-9 


In malarial fever, therefore, there is a marked shift to the left of 
the Ameth count, which may be extreme in degree in cases accom¬ 
panied by severe constitutional symptoms, but is still quite definite 









448 


even in cases in which there is no proof of the nature of the infection 
other than the clinical and therapeutic one. In severe cases the 
majority of the polymorphonuclear leucocytes are of the types 
included in Class I, and forms with simple horseshoe-shaped nuclei 
are peculiarly abundant. On examining a blood film from such a 
case one is at once struck by the uniform and unfamiliar appearance 
of the polymorphonuclear leucocytes, and the almost complete 
absence of the familiar normal forms with the nucleus divided into a 
number of separate lobes joined together by delicate threads. 

This effect on the neutrophile leucocytes is observable before the 
onset of the attack of malaria, and before the parasites are 
sufficiently numerous to be detected in an examination of the 
peripheral blood. An example may perhaps be given. On 
April 2 ist I examined the blood of a European who had landed at 
Accra three weeks previously. He had recently returned from 
leave, and had not taken any quinine for six months, but had 
enjoyed good health during all this time. I found that there was a 
marked shift to the left of the Ameth count, and that the index was 
61*5; and although no parasites and no pigmented leucocytes could 
be found, and the percentage of the large mononuclear cells was not 
higher than that frequently found in healthy Europeans in West 
Africa, I suggested that it was probable that he had been recently 
infected with malaria. A few days later he developed a typical 
attack of malarial fever. 

This shift to the left may persist for a considerable time after the 
attack of malaria is apparently cured. In Table VI the results of 
five successive Ameth counts on the same individual are shown. 


Table VI.—Successive Ameth counts in a severe case of malarial fever in a native. 


Date 

Clinical condition 

Arneth classification per cent. 

' 

Ameth 

index 

I 

1 II 

m 

IV | 

V 

Jan. 

14, 1915... 

P, falciparum , 1 to 400 R.B.C.... 

67*0 

! 25-0 

8-o j 

— | 

— 

92*0 

Jan. 

15, 1915... 

P. falciparum, l to 713 R.B.C.... 

46*0 

43 ’° 

11*0 

— I 

— 

89*0 

Jan. 

25,1915... 

Well . 

2 7*5 

47-0 

22*0 

3*5 

— 

74*5 

Feb. 

9,1915... 

Well . 

20*0 

4*’5 

| 

34*5 

1 

3 -o 

— 

62-5 

Feb. 

2+, 1915... 

Well . 

18.5 

43*5 

34-0 

4 -o j 

— 

62*0 




+49 


This patient, a native, was treated with quinine only during the 
actual attack of fever; and although he made a rapid and apparently 
complete recovery, there was still a decided deflexion of the Arneth 
count six weeks after the onset of his illness. 

In West Africa at the present time repeated infection with the 
parasites of malarial fever must be recognised as one of the normal 
factors of everyday life alike for the native and the European. The 
native by his tolerance, and the European by his prophylactic doses 
of quinine, is able, as a rule, to ward off the actual attack of fever; 
but the parasites are continually being introduced into the system, 
and sooner or later they obtain a foothold. Once infected, it is well 
known how difficult it is to eradicate them completely from the body. 
Bearing these facts in mind, I think it is only natural that the 
average native and European in West Africa should show evidences 
of malarial infection, and I believe that the shift to the left of the 
Arneth counts found in healthy individuals (see Section I) are to be 
explained in this manner. 

III. THE RESULTS OF ARNETH COUNTS IN YELLOW FEVER 

The early diagnosis of yellow fever is a matter of great difficulty, 
especially in the mild attacks which occur in natives in West 
Africa. In the Second Report of the Yellow Fever Commission 
(West Africa), one of the conclusions arrived at is that * the extreme 
practical importance of being able to determine whether a mild case 
of fever is or is not yellow fever renders it essential that all possible 
methods should continue to be employed in the clinical study of the 
disease/ and it is recommended that ‘ the attention of all workers at 
this subject should be specially directed to the discovery of a 
clinical test for yellow fever/ 

I was anxious, therefore, to carry out Arneth counts in cases of 
yellow fever, since this test, should it prove to be of any assistance, 
would be a very simple one for the physician to employ, as it does 
not entail any elaborate technique. Unfortunately, the single case 
of this disease that has occurred at Accra since my arrival was only 
identified at the autopsy. Recently, however, Dr. J. M. O’Brien 
has been so kind as to hand over to me a series of blood-films he 
had collected from cases of yellow fever at Guayaquil, Ecuador; 
and has permitted me to make use of them for the purposes of this 
investigation. I wish to take this opportunity of thanking 



450 

Dr. O'Brien for his generosity in allowing me to make use of this 
material. 

The slides were from the cases of yellow fever in which 
Dr. O’Brien (1914) detected a degeneration of the polymorpho¬ 
nuclear leucocytes which he believes to be characteristic of the 
disease. On examining them I found that they had been taken on 
thirty-five days from seventeen different cases of yellow fever. In 
seven cases there was only a single blood-film, but in the other ten 
there were two or more, taken on successive days of the disease. 
Ameth counts have been made on all the thirty-five films, but I 
shall limit myself for the present to a consideration of those made 
on the first days of observation only, the details of the counts on 
which will be found in Table VII. 


Table VII.—Ameth counts in yellow fever. 


No. 


Arneth classification f 

>er cent. 


1 

! Arneth 


I 

II 

III 

IV 

V 

index 

1 

1 

43-0 

41*0 

1 5*5 

1 o -5 

— 

■ 

0 

rh 

00 

2 

48*0 

‘ 40-5 

9-5 

1 2-0 

— 

I 88-5 

3 

67-5 

30-0 

2*0 

°*5 

— 

97*5 

4 

69*0 

30*0 

1*0 

— 

— 

99.0 

5 

82*0 

17*0 

1*0 

— 

— 

99 ‘° 

6 

40*0 

40*0 

19*0 

1*0 

— 

8o*o 

7 

49-0 

44.0 

7 *o 

— 

— 

93 *o 

1 

8 

58-5 

36*5 

5 -o 

— 

— 

1 95-0 

| 

9 

75 *o 

24*0 

1*0 

! 

— 

99 *o 

10 

34-0 

50*0 

15*0 

1*0 

— 

00 

4* 

6 

11 

53 *o 

43*0 

4 -o 

— 

i 

96-0 

12 

20*0 

5 2 *5 

22*0 

5 -o 

°*5 

7 2 -5 

>3 

55*5 

34 *o 

8*5 

2*0 

_ i 

89-5 

*4 

59*5 

36-5 

4 *o 

— 

__ 1 

96*0 

*5 

49.0 

41*0 

10*0 

— 

~ | 

90-0 

16 

42-5 

36*0 

20*5 

1*0 

__ 1 

1 

785 

<7 

6o-o 

37‘5 

2*0 

o-s 


97-5 



37-16 

8*64 

079 

0*02 1 

l 

90*52 



451 


The examination showed that there was a profound shift to the 
left of the Arneth count in yellow fever. The index on the first days 
represented in the collection of blood-films averaged 90* 5, and in 
some cases actually 99 per cent, of the polymorphonuclear leucocytes 
belonged to Classes I and II. Even after allowing liberally for a 
possible personal factor in the counts, such a deflexion is much 
greater than has been described by other observers in any other 
disease. As in the severer cases of malaria, a great many of the 
cells had a simple horseshoe-shaped nucleus, and in one patient 
these forms constituted 82 per cent, of the polymorphs. 

The shift to the left of the Arneth count in yellow fever is more 
pronounced than it is in malaria. An attempt to show this fact 
graphically has been made in Charts III and IV. Chart III shows 
the average Arneth counts in yellow fever, in malaria, and in normal 
persons in Europe. Chart IV shows the curves produced by plotting 
the Arneth indices by percentages into ten equal groups in cases of 
yellow fever, in Europeans suffering from malaria, and in 
apparently healthy Europeans in Accra. 

It has been shown above that in the twenty-three cases of malaria 
studied at Accra in which the diagnosis was confirmed by the 
finding of the parasites, the Arneth index averaged 86*6 as 
compared with 90*5 in these seventeen cases of yellow fever. The 
difference is one of degree; and as in some of the cases of yellow 
fever the index was lower than it was in some of the cases of 
malaria, the index alone could not be used as a means of making a 
differential diagnosis between these two diseases from an examina¬ 
tion of the blood on the first day on which a case happened to be 
seen. 

In those cases of malaria, however, in which the evidence of 
infection was presumptive only, the average index was found to be 
79*3; and in the cases of fever merely suspected of being malarial, 
it was lower still, namely, 71 ’35. It is in cases of the latter type 
that it is important to be able to decide whether or not they are 
yellow fever. The occurrence of a very high Arneth index in any 
suspicious case of fever in which no malaria parasites could be 
found, and in which there was not presumptive evidence of malaria, 
would, I believe, be strongly in favour of a diagnosis of yellow 
fever. It will be necessary, before arriving at a definite decision, to 



452 


study the condition in mild cases in natives, and to follow the 
changes in the Ameth counts from the earliest stages of the disease 
until convalescence is fully established. I hope to be able to do this 
as soon as I can procure sufficient local material. 



Chart III.—The average Arneth counts in (A) normal persons in 
Europe (Arneth), (B) twenty-three cases of malarial fever at 
Accra, and (Q seventeen cases of yellow fever 





453 



Chart IV.—An attempt to show graphically the degree of the shift to 
the left in the Arneth counts in (A) healthy Europeans, (B) Euro¬ 
peans suffering from malaria, and (C) yellow fever. 










454 


IV. THE SIGNIFICANCE OF THE CHANGES OBSERVED IN THE 

ARNETH COUNTS 

A marked shift to the left of the Arneth count has generally 
been assumed to be an indication of a lowered resistance. As the 
result of a large number of examinations in various diseases, Arneth 
concluded that there was a definite relationship between the blood 
picture and the progress of the disease; and the view adopted by 
him, and by most subsequent authors, was that the cells of Classes 
I and II were younger, and less efficient than the cells of the higher 
classes; and that therefore the degree of the shift to the left was an 
index of the patient’s resistance. Chamberlain (1914) summarises 
this view by saying ‘ a high Arneth index (excess of Classes I and 
II) goes hand in hand with a low resistance, or with a high degree 
of toxic and bacterial absorption which is bringing about the 
destruction of the actively phagocytic leucocytes (Classes III and 
IV).’ 

There is, however, little evidence in support of the assumption 
that the neutrophile leucocytes of Classes I and II are less actively 
phagocytic than those of the higher classes. Many observers have 
failed to detect any difference in experiments carried out in vitro; 
and I have recorded elsewhere a case of fatal malarial fever in which 
the leucocytes had ingested a large number of parasites, making it 
possible to gauge their phagocytic activity in vivo> and have shown 
that in this case the polymorphonuclear leucocytes of Class I were 
certainly not less efficient than those of the higher classes. 

Breinl and Priestley (1914) have, therefore, advanced the view 
that ‘ the Arneth picture is an expression of the functional activity 
of the leucopoetic system, especially the bone-marrow, rather than 
that of phagocytic activity.’ They dissent from the view of 
Chamberlain and Vedder that the shift to the left observed in 
Filipinos may be an indication of a lowered resistance to infections 
on the part of the native races, and attribute the similar phenomenon 
found in North Queensland children to purely climatic influences. 
They state that in their experience there is amongst the children in 
Townsville, who show a decided shift to the left of the Arneth count, 

‘ no greater susceptibility to infectious diseases than amongst the 
same class in Europe.’ 



+55 


In West Africa the healing powers of the natives are famous, 
and it is scarcely credible that the slight shift to the left of the 
Ameth count observed in them can be an indication of a lowered 
resistance to infection. As I have already stated, I believe that in 
West Africa the shift in apparently healthy individuals may be 
accounted for by infections with malaria parasites which do not 
necessarily culminate in an attack of 'fever.* 

The very pronounced shift to the left observed in confirmed cases 
of malarial fever at Accra may be considered as supporting the view 
of Breinl and Priestley that the phenomenon is an indication of the 
functional activity of the leucopoetic system; for on this hypothesis 
it is in such diseases as malaria, which profoundly affect the spleen 
and the bone-marrow, that one might expect to find a marked shift 
to the left indicating a morbid activity of these tissues. The Arneth 
index was highest in those cases which showed the greatest 
constitutional disturbances, and not necessarily in those that had 
the largest number of parasites in the peripheral blood. In yellow 
fever, another disease accompanied by severe constitutional 
disturbances and a profound toxaemia, the Arneth counts showed a 
shift to the left that was often extreme in degree. 

The blood-films from yellow fever cases that were examined, were 
some of those from which 0 *Brien (1914) described a degeneration 
of the polymorphonuclear leucocytes which he considered to be 
characteristic of the disease. Some of the abnormal forms he 
interpreted, correctly I believe, as being dead cells. Identical forms 
were described by Dr. Mary Rowley (1907) in a fatal case of aortic 
and mitral disease with anaemia; and this author was able to bring 
forward evidence in proof of the fact that these cells had been killed. 
Similar polymorphonuclear leucocytes are seen occasionally in the 
blood of normal persons; but they are relatively much more 
abundant in acute cases of malaria and in yellow fever. It is 
probable that in these diseases the profound toxaemia causes a great 
destruction of the circulating polymorphonuclear leucocytes, and 
thus leads to a relative increase in the peripheral blood of the young 
cells—namely, those belonging to Ameth*s Classes I and II, and 
a corresponding shift to the left of the count. 

The increase in the percentage of young cells in the peripheral 
blood might result from (1) the elimination of the cells of the higher 



456 


classes by the selective action of the toxins as suggested by 
Chamberlain, or (2) a flooding of the blood with newly-formed 
elements produced by the activity of the leucopoetic system. As a 
leucocytosis is frequently observed in blood in which there is a 
marked shift to the left of the Ameth count, and as there is really 
no reason to suppose that the cells of the higher classes are the more 
actively phagocytic, and so the more liable to be destroyed by a 
high degree of toxic and bacterial absorption, I am inclined to think 
that the latter explanation is the correct one. 

It might be expected, then, that in an intense infection there would 
be developed an immediate shift to the left of the Ameth count, 
increasing rapidly in degree as the toxaemia deepened, and tending 
to diminish as recovery took place. 

SUMMARY 

1. A slight shift to the left of the Arneth count is found in the 
blood of healthy Europeans in West Africa. 

2. There is a marked shift to the left in malarial fever which is 
evident not only during the attack, but also before its onset, and 
for a considerable time after convalescence is established. 

3. It is probable that the abortive inoculations with malaria 
parasites by infected mosquitos, which are a part of the daily life in 
many parts of West Africa, are sufficient to account for this shift in 
apparently healthy Europeans, without postulating a specific action 
of the climate on the white races living in the tropics. 

4. In yellow fever there is a great shift to the left of the Arneth 
count. 

5. It is suggested that the changes observed in the Ameth 
counts are due to toxaemia causing a destruction of the circulating 
polymorphonuclear leucocytes, and a flooding of the blood with 
young cells liberated by the activity of the leucopoetic system. 

REFERENCES 

Buinl, A., and Priestley, H. (1914). Annals of Trop. Med. and Parasitol., Vol. VIII, p. 565. 

Chamberlain, W. P. (1914). American Jour, of Trop. Diseases, Vol. II, p 48. 

Cooke, W. E. (1915). Jour, of Path, and Bact., Vol. XIX, p. 494. 

Knapp, H. H. G. (1915). Indian Medical Gazette, Vol. L, p. 95. 

Macfie, J. W. Scott (1915). Lancet, May 1. 

Rowley, Mary W. (1907). Jour, of Experim. Med., Vol. X. p. 78. 



457 


BABESIASIS AND TRYPANOSOMIASIS AT 
ACCRA, GOLD COAST, WEST AFRICA 

BY 

J. W. SCOTT MACFIE, D.Sc., M.B. 

WIST AFRICAN MEDICAL STAFF 

(Received for publication 4 August , 1915) 

Plates XXXV, XXXVI 

CONTENTS 

I. Babesiasis. page 

A. Babesiasis of cattle and sheep . 457 

B. Canine Babesiasis . 462 

C. A piroplasm, NuttdUia decumani , n.sp., of the brown rat (Mus dtcumanus ) 462 

II. Trypanosomiasis. 

A. Trypanosomes found in blood films obtained from the Accra slaughter¬ 

house ... 464 

B. Canine Trypanosomiasis . 477 

C. Equine Trypanosomiasis . 478 

D. A small monomorphic Trypanosome found in the blood of a mare ... 480 

E. Two cases of Trypanosomiasis in mules resembling acute Do urine ... 486 


BABESIASIS 

A. BABESIASIS OF CATTLE AND SHEEP 

In a previous paper (1914) I gave a preliminary account of the 
occurrence of babesiasis in Nigeria, and described three species of 
parasites found in the blood of cattle and sheep in that country. 
Similar observations have been made by Bouet (1908) on the Ivory 
Coast, and by Broden and Rodhain (1909) in the Congo; the former 
recording the occurrence of P. parvum, P. mutans and P. bigeminum, 
and the latter P. mutans and P. bigeminum. From these observa¬ 
tions it is evident that piroplasms are common on the West Coast, 
and it may therefore be of some interest to record for comparison the 
results of the examinations of domestic animals at Accra, in the Gold 
Coast, since hitherto but little attention has been directed to these 
parasites in British West Africa. 

For the purposes of this investigation blood films were obtained 



45 8 


from the Accra slaughter-house. Hump-backed cattle, straight- 
backed cattle, sheep, pigs, and goats are slaughtered daily at Accra. 
Many of the animals are bred locally, but others come from a 
distance, and no doubt bring with them their parasitic infections. 
The hump-backed cattle, as in Nigeria, are bred in the north, and 
are driven south in herds during the dry season until they reach the 
coast towns. On the journey they are exposed to the attacks of ticks 
and tsetse-flies, and debilitated by fatigue, they are probably but 
feebly resistant to parasitic infections. The straight-backed cattle, 
on the other hand, are mostly bred in the southern parts of the Gold 
Coast, and the majority of those examined were either Accra or 
Addah animals; they had not, at any rate, treked down from the 
north like the hump-backed cattle. A considerable number of the 
sheep had also come to Accra from a distance; but the 
majority of the goats, and probably all the pigs, had been bred in 
the neighbourhood. 

A hundred animals of each kind were examined, namely hump¬ 
backed cattle, straight-backed cattle, sheep, pigs, and goats. Only 
a single blood film was obtained from each animal, so that the 
percentages of infections found are probably considerably below 
those actually occurring. In this way piroplasms were found in 
53 per cent, of the hump-backcd cattle, in 40 per cent, of the straight- 
backed cattle, and in 21 per cent, of the sheep (Table I). None were 
found in the pigs and goats. 


Table I.—The results of the examination of 500 domestic animals for babesiasis. 


Host 

Number 

examined 

Number 
, infected 
with 

Piroplasms 

B. bigemina 

T. mutans 

Cattle, hump-backed breed . 

100 

1 

53 

7 

49 

Cattle, straight-backed breed . 

100 

40 

3 

38 

Sheep . 

100 

21 

— 

21 

Pigs. 

100 

— 

— 

— 

Goats . 

100 

— 


— 

Totals. 

500 

114 

10 

108 





459 


So far as could be ascertained the infections appeared to be 
benign, but as the animals had frequently concurrent infections with 
other parasites, especially trypanosomes, it was difficult to determine 
this point. For instance, forty-seven of the fifty-three hump-backed 
cattle found to harbour piroplasms were infected with trypanosomes 
also. In the case of the sheep no trypanosomes were found 
associated with the piroplasms, and in these animals no clinical 
symptoms of disease were observed. 

Various abnormal conditions of the red corpuscles were found in 
the blood films, notably a coarse basophilia. Anaplasma-like bodies 
were present in many cases, but as these are common in healthy 
animals they can be of no special significance in this connexion. In 
the blood films from the cattle, especially the straight-backed breeds, 
there were generally to be found oval or rounded bodies about the 
size of a red blood corpuscle that stained a pale blue colour. A few 
granules that stained similarly to chromatin were present in these 
bodies. These basophile cells were similar to those described by 
Castellani (1912) as occurring in man in cases of yaws, psoriasis, 
lichen, acne, severe anaemia, &c.; and they were probably, as he 
concludes, 4 red cells undergoing degenerative changes/ Although 
piroplasms were present in the blood of most of the cattle in which 
they were detected, there does not appear to be any direct connexion 
between them and these parasites, since they were not constantly 
associated. The basophile cells may, however, have been an 
indication of the anaemia that is frequently induced by piro- 
plasmosis, although, as has already been stated, the infections 
appeared otherwise to be benign. 

Two species of piroplasm were encountered in the blood films, the 
one a large pyriform parasite, and the other a small oval or rod¬ 
shaped organism. Both types were found in cattle, and the latter 
in sheep also. The third species found in Nigeria, a large parasite 
with forms in which the chromatin was divided into six or seven 
masses, was not found in the Gold Coast. 

Owing to the lack of unquestionably uninfected animals for 
experiments, it has not been possible to attempt to determine the 
ticks that transmit these parasites. Experiments designed to decide 
this point could not be conclusive if carried out with locally bred 
animals in the Gold Coast because of the high percentage of natural 



460 


infections in the cattle and sheep, the benign nature of the infections, 
and the extreme rarity of the parasites in some cases. Some ticks 
collected from the animals were, however, sent home to England, and 
were very kindly identified by Prof. Nuttall. The species from both 
cattle and sheep were the same, namely, Boophilus y Ambylomma 
variegatum , and Hyalomma aegyptium; and presumably one or 
more of these transmits the piroplasms. 

Babesia bigemina 

The larger species of piroplasm appeared in the blood as oval or 
pear-shaped bodies of considerable size. Pairs of pyriform bodies 
were not uncommon, and when found were generally seen to occupy 
the greater part of the enveloping erythrocyte. There can be little 
doubt that these parasites should be identified as Babesia bovis or 
bigemina (Piroplasma bigeminum). In the straight-backed cattle 
the morphology of the parasites was similar to that found in dwarf 
cattle in Nigeria (1914); but in the hump-backed cattle the pyriform 
pairs were somewhat smaller, and their nuclear structure was less 
distinct. In one hump-backed beast a considerable number of the 
parasites were notably amoeboid. 

This type of Babesia was found in 7 per cent, of the hump¬ 
backed cattle, and in 3 per cent, of the straight-backed breed. 
They were never very numerous, and in four of the ten animals in 
whose blood they were detected they were associated with the smaller 
type of parasite to be described immediately. 

Theileria mutans 

The smaller type of piroplasm was identical with that found in 
Nigeria. At Accra it was present in 49 per cent, of the hump-backed 
cattle, in 38 per cent, of the straight-backed cattle, and in 21 per 
cent, of the sheep. It was, therefore, a very common parasite of the 
animals brought to the Accra slaughter-house, just as it was of the 
domestic animals in Nigeria. In several of the hump-backed cattle, 
and a few of the sheep, the infection was a heavy one; but in the 
straight-backed cattle the parasites were as a rule scanty or rare. 
Certainly in the majority of the animals the infection appeared to 
be benign. 

The parasites were very simple but remarkably pleomorphic 
organisms, appearing as ring-shaped, oval, horseshoe-shaped, and 



461 


rod-like bodies, and characteristic cross-forms. Examples of all 
these different forms were figured in my account of the Nigerian 
infections, so that it will be unnecessary to illustrate the present 
description. Since then I have had opportunities of examining 
smears from the spleen in cases of several infected animals, but up 
to the present I have not detected the presence of Koch's 'blue 
bodies' in any of them. Such a parasite, according to the classifica¬ 
tion of Franca, would have to be included in the genus Theileria , 
and for this reason, in describing its occurrence in Nigeria, I 
provisionally identified it as Theileria parva > stating at the same 
time that Koch's 'blue bodies' had not been found, and that the 
infections appeared to be benign. Minchin (1912), however, 
considers that ‘ the diagnosis of the genus Theileria given by Franca 
would appear to apply to B. mutans rather than to T. parva ,' and 
states that 'a confusion has arisen between two parasites very 
similar as regards the appearances they present in the blood, but 
differing in every other respect, namely, Theileria parva , the true 
parasite of " East Coast fever " of cattle, and Babesia (Piroplasma) 
mutans , also found in cattle. In both parasites alike the charac¬ 
teristic cross-forms appear in the blood. In Theileria parva , 
however, the cross-forms are an aggregation of four distinct 
gametocytes which have invaded the same corpuscle, while in 
Babesia mutans the cross-forms are produced by quadruple fission 
of an ordinary multiplicative individual.' The parasite described 
from Nigeria, and that found recently in the Gold Coast, showed 
cross-forms unquestionably produced by fission, and not by the 
aggregation of four distinct gametocytes in a single cell. It should 
therefore be identified, according to Minchin, as B. mutans. The 
facts that the infections appear to be benign, and that Koch's ' blue 
bodies' have not hitherto been detected in smears from the spleen 
of infected animals, tend also to prove that the parasite is not 
T. parva. Inoculation experiments would assist the diagnosis, as 
T. parva is not inoculable; but for the reasons given above these 
could not be conclusive if carried out in this country. At the same 
time, an organism characterised by the occurrence of bacilliform or 
rod-shaped parasites, and multiplicative forms in the shape of a 
cross, cannot well be assigned to the genus Babesia , and it will 
probably be best for the sake of clearness to identify this small 
piroplasm as Theileria ( Babesia ) mutans. 



462 

B. CANINE BABESIASIS 

Up to the present only a single case of canine babesiasis has 
come under my notice at Accra. The dog found to be infected had 
been brought into the Colony only six weeks previously. He was 
feverish, much wasted, and very anaemic. The right eye was 
dimmed, and there was a profuse watery discharge from the nose. 
On examining the blood, large numbers of piroplasms were found; 
many extremely amoeboid, but others showing the pyriform outline 
and the association in pairs characteristic of Babesia cants. 

Imported dogs are known to suffer on the Gold Coast from a 
disease characterised by severe anaemia and rapid emaciation which 
does not appear to be a trypanosomiasis. Possibly some of the cases 
of this disease are due to babesiasis. It is the common experience 
that in Accra dogs become infested with ticks, no matter how 
carefully they are tended, and it would not therefore be surprising 
if babesiasis were found to be a prevalent disease. 

No case of equine babesiasis has as yet come under my notice 
in the Gold Coast, although this disease is known to occur in 
Nigeria and in the neighbouring colonies. 

C. A PIROPLASM, NUTT ALU A DECUMANI, n. sp., OF THE BROWN 
RAT (MUS DECUMANUS ). PI. XXXVI, figs. 1-14 

In the blood of a few brown rats {Mus decumanus) that had 
either been sent to me for examination by the Medical Officer of 
Health for Accra, or had been caught in the laboratory itself, 
unpigmented parasites were found in the erythrocytes. Four out 
of twenty rats recently examined were found to be infected; that is, 
a proportion equal to 20 per cent. 

From two of the rats only a single blood-film was obtained, and 
in both of these the parasites were rare. In the other two the 
infections were heavier, and as I was able to keep the animals in 
captivity, it was possible to make a more thorough study of the 
morphology of the parasites. Both these rats were young animals, 
and both showed in their red corpuscles, in addition to the piro¬ 
plasms, the bodies known as GrahameUa. The first was caught on 
January 10th, 1915, and was kept under observation until February 
15th, the thirty-seventh day, when it was accidentally killed. When 
caught, the examination of the blood showed the presence of 



4*3 


Grahamella and Jolly bodies, but no piroplasms. A few days later 
one or two piroplasms were detected, and by the fifteenth day a 
considerable number were present, although the infection was never 
a heavy one, and the parasites had always to be searched for most 
carefully. From this date the parasites gradually diminished in 
numbers, so that by the twenty-sixth day they were rare. They 
were still present on the thirty-seventh day when the rat was killed. 
The Grahamella bodies were not found after the thirteenth day. 
The second rat was caught on April 27th, and is still under 
observation. Piroplasms were found in the blood on the first day, 
and continued to be present up to the tenth day, but have not been 
detected since. They were always rare, and it was only after 
prolonged search that they were found. The Grahamella bodies 
have vanished from the blood of this rat also since it has been under 
observation. 

The forms of the parasite most commonly seen consisted of a 
somewhat irregularly shaped chromatin mass, and an amoeboid or 
ring-shaped protoplasmic body. The smallest parasite seen was a 
minute dot of chromatin to which a very small ring of blue-stained 
material appeared to be attached, but even at this early stage of 
development there was a paler area or vacuole in the protoplasm 
(PI. XXXVI, figs. 1 and 5). More mature parasites were often of 
considerable size, and great irregularity of outline. The chromatin 
was in many cases drawn out into a number of processes or separated 
into two or more distinct masses, and the protoplasm was freely 
vacuolated or extended in a number of delicate threads. Other 
parasites were signet-ring-shaped, and closely resembled the 
characteristic forms of Plasmodium falciparum . Sometimes there 
were two chromatin dots at opposite sides of the ring. None of the 
parasites was pigmented. The red corpuscles did not appear to be 
affected by the presence of the parasites, and were neither obviously 
enlarged nor stippled. 

After long search a few specimens were found in which division 
had taken place, resulting in the production of four lanceolate 
parasites arranged in the form of a cross. Each of these bodies 
contained a nucleus which was generally situated near the middle, 
and in addition a second minute chromatin dot could usually be 
seen at the distal end. No pairs of pyriform bodies were found. 

Subcutaneous inoculations of blood containing these parasites 



+64 


were made into three white rats, but without transmitting the 
infection. A single attempt to convey the parasites to another brown 
rat, by means of ticks taken from the body of an infected animal, 
was also unsuccessful. 

This parasite would seem to belong to the genus Nuttallia , and 
as I am unable to find any reference to a previous description of it, 
I would suggest the name Nuttallia decumani for it, should it prove 
to be a new species. 


REFERENCES 

Bourr, G. (1908). Piroplasmoie bovine observte A la c6te d’Ivoire. Bull . Soc. Path. Exot., 

Vol. I, p. 234. 

Broden, A. and Rodhain, G. (1909). Piroplasmoses des bo vide* observees au Stanley-Pool. 
Bull. Soc. Path. Exot., VoL II, p. 120. 

Castellani, A. (1912). Note on Certain Cell Inclusions. Journ. Trop. Med. and Hyg., Vol. 
XV, p. 354. 

Macfie, J. W. Scott (1914). Notes on some Blood Parasites collected in Nigeria. Annals 
of Trop. Med. and Parasit., Vol. VIII, p. 439. 

Minchin, E. A. (1912). An Introduction to the Study of the Protozoa, p. 380. Edward 
Arnold, London. 


TRYPANOSOMIASIS 

Three types of trypanosomes were met with at Accra, namely, 
(1) a polymorphic parasite, (2) a large monomorphic parasite with 
a long free flagellum, and (3) a short monomorphic parasite without 
a free flagellum. These three types are the same as those found in 
Nigeria, and which I have elsewhere identified as (1) T. fecaudi 
(T. brucei of Uganda, T. ugandae ), (2) T. vivax , and (3) T. pecorum 
( T. congoletise). In dried blood-films, such as those obtained from 
the slaughter-house, morphological features had to be relied on for 
the identification of the trypanosomes. Such data are, of course, 
insufficient to distinguish the more closely allied species; but as 
the small number of inoculations into animals that were made seemed 
to confirm these diagnoses, they are probably correct. 

A. TRYPANOSOMES FOUND IN BLOOD FILMS OBTAINED 
FROM THE ACCRA SLAUGHTER-HOUSE 

Blood-films were examined from 500 animals slaughtered at 
Accra, namely, 100 each from hump-backed cattle, straight-backed 
cattle, sheep, pigs, and goats. The films were the same as those 



+65 


used in the examinations for Babesiasis, of which an account has 
already been given; and the remarks made there with regard to the 
sources from whence the animals came should be taken into considera¬ 
tion with reference to the trypanosome infections. In Accra itself 
tsetse flies are exceedingly rare. A few are caught every year, but 
it is generally supposed that these have found their way into the 
town in railway carriages or waggons. Within a few miles, 
however, tsetse flies are numerous; and any animals coming from 
a distance must be exposed to the attacks of these insects for the 
greater part of their journey. It is probable, therefore, that the 
majority of the trypanosome infections described had been con¬ 
tracted before reaching Accra. 

Hump-backed Cattle. Trypanosomes were found in 92 per 
cent, of the hump-backed cattle; and as only a single blood-film 
was examined from each animal, this proportion is more probably 
below the mark than above it. In the majority the parasites were 
very numerous. Trypanosomes of the type of T . vivax were present 
in 76 per cent., T . congolense in 28 per cent., and T. pecaudi in 
12 per cent, (see Table II). Sixty-nine of the hundred animals 
examined were infected with a single type of trypanosome, fifteen 
with T. vivax and T. congolense , four with T. vivax and T. pecaudi , 
three with T. congolense and T. pecaudi , and one was the host of 
all three types, T. vivax , T. congolense , and T. pecaudi . 

Tablx II.—Trypanosdme infections found in animals killed at the Accra slaughter-house. 



Number 

Number 1 Percentages infected with 

infected | 

Host 

examined 

with i 

trypanosomes T. vivax 

T. congolense 

T. pecaudi 


Cattle, hump-backed 

i 

100 

9 Z 

76 

j 28 

! 1 2 

Cattle, straight-backed 

>00 ! 

18 

14 

6 

1 

Sheep . 

100 

4 

3 

2 

- 

Pigs. 

100 

5 

— 

5 

— 

Goats 

100 

1 

1 

1 


Totals 

500 

120 

• iX-8 

8-4 

2-6 




466 


Of all the animals slaughtered at Accra the hump-backed cattle 
have the longest journey to come, and this fact probably accounts 
for the great number of them found to harbour trypanosomes. Even 
as far north as Kumasi, I understand, almost all are found to be 
already infected with these parasites. 

Straight-backed Cattle. Trypanosomes were found in 18 per 
cent, of the straight-backed cattle. The parasites were generally 
scanty, and were often detected only after a long search. T. vivax 
was found in 14 per cent., T . congolense in 6 per cent., and 
T . fecaudi in 1 per cent. Fifteen of the animals had a single 
infection, and three harboured both T. vivax and T. congolense . 

Straight-backed cattle are bred in the southern parts of the Gold 
Coast, and many of those examined at Accra had not been brought 
from any great distance. The relatively low percentage of trypano¬ 
some infections, and the rarity of the parasites in the blood, should, 
probably, be correlated with this fact. Some of these animals were, 
however, of the dwarf breed peculiar to West Africa, and it should 
be remembered that, as I have suggested elsewhere (1913), they may 
possesses a partial immunity to T . vivax at any rate. 

SHEEP. Four of the hundred sheep examined were found to be 
infected with trypanosomes, but in each case the parasites were rare. 
T. vivax was found in 3 per cent., and T. congolense in 2 per cent. 
Three sheep had a single infection, and one harboured both species 
of trypanosome. 

PlGS. Five out of the hundred pigs examined were found to be 
infected with trypanosomes, and in each case the type of parasite 
was T. congolense .* The parasites were generally very rare. 

GOATS. Only one of the goats was found to harbour trypano¬ 
somes, and this animal had a double infection— T. vivax and 
T. congolense. It is somewhat remarkable that so few infections 
should have been found in goats, since these animals have been 
credited with being a natural reservoir for T. vivax , the trypanosome 
most frequently met with at Accra. 

T. pecaudi ( T . brucei of Uganda, T. ugandae) 

Polymorphic trypanosomes were found in thirteen animals 
(2 6 per cent.), namely, in twelve hump-backed cattle, and in one 

• Subsequently a pig was examined in who?e blood trypanosomes of the T. pecaudi type were 
present. 



4 6 7 


straight-backed cow. This type of trypanosome was the most 
uncommon of the three forms occurring at Accra. It is the most 
fatal to domestic animals in Nigeria, and so probably in the Gold 
Coast also. In the hump-backed cattle it was often associated with 
other trypanosomes: four times with T. vtvax , four times with 
T. congolense , and once with both T. vtvax and T. congolense. 

Morphologically the trypanosomes resembled T. pecaudi or 
T. brucet of Uganda, both long and slender, stumpy, and inter¬ 
mediate forms were found in each case, but forms with posterior 
nuclei were not seen. It is impossible, therefore, to exclude the 
possibility that some of the trypanosomes might have been 
T, gambiense. The occurrence of posterior nuclear forms is, 
however, very irregular in cattle infected with trypanosomes which, 
when subinoculated into rats and guinea-pigs, show high percentages 
of these forms; and at the periods during which the parasites are 
scanty, as they were in all the slaughter-house cases, it is generally 
impossible to find them at all. Too much importance should not, 
therefore, be attached to the absence of these forms, especially when 
the materials examined are restricted to dried blood-films. 

In its behaviour in the tsetse fly, T. pecaudi is said to differ 
from the other polymorphic trypanosomes, development taking place 
in the gut and proboscis, instead of in the gut and the salivary 
glands. Confirmation of this observation is desirable; but in any 
case, in dealing with these slaughter-house films the exogenous cycle 
of development had to be ignored, and it was impossible to carry 
identification beyond the point at which the trypanosomes were 
assigned to what is sometimes known as the ‘ T . brucei group.* 

In one instance, blood containing this type of trypanosome was 
obtained from a hump-backed ox, and inoculated subcutaneously 
into a white rat and a guinea-pig. The rat first showed trypano¬ 
somes in its blood on the tenth day, and death took place on the 
nineteenth day. The parasites were at first very scanty, but 
gradually increased in numbers up to the day of death; and in the 
latter stages of the disease, forms with posteriorly placed nuclei were 
numerous. The guinea-pig did not become infected, but with two 
others was successfully inoculated with blood taken from the rat. 
In these three guinea-pigs the incubation periods were respectively 
twenty, nineteen, and seven days; and the duration of the disease 
thirty-four, twenty-five, and thirteen days. 



468 


Measurements were made of 200 trypanosomes from the blood of 
the rat (see Table III, and Chart I). The average length was 
27 66 p; the longest individual measuring 39 p, and the shortest 
15 M- This trypanosome then corresponded with T. pecaudi 
(T. brucei of Uganda), both in its morphology and in its patho¬ 
genicity to rats and guinea-pigs. 


Table III. —Measurements in length of T. pecaudi (cattle strain). 


Animai 


Day of 
the 

disease 


Number 

measured 


Length in microns 


Average Minimum I Maximum 


! 


Rat . 

1 

2° 

Rat 

2 

2 5 

Rat . 

3 

20 

Rat . 

4 

2 5 

Rat . 

5 

. 20 

Rat . 

6 1 

2 5 

1 

Rat . 

7 

20 

Rat . 

8 

2 5 

Rat . 

9 

20 


'i 

200 


*7-65 

16 

33 

2 77 2 

! «7 

37 

26*25 

16 

35 

3 2 *64 

26 

| 

39 

26*65 

1 

18 

33 

26*88 

16 

35 

28*2 

16 

38 

27-2 

‘5 

39 

24-8 

16 

35 

27*66 

*5 

39 










4<*9 


















470 


T. vivax 

Trypanosomes morphologically identical with T. vivax were 
present in ninety-four (18*8 per cent.) of the animals examined, 
namely, in seventy-six hump-backed cattle, fourteen straight-backed 
cattle, three sheep, and one goat. They caused by far the greatest 
number of the trypanosome infections met with at Accra, and in 
many of the animals, especially the hump-backed cattle, were 
present in the blood in enormous numbers. 

Blood from hump-backed cattle heavily infected with these 
trypanosomes was inoculated into two white rats, two guinea-pigs, 
and one rabbit, but without result. The insusceptibility of the 
smaller laboratory animals, taken in conjunction with the very 
characteristic morphology of the parasites, is sufficient evidence for 
the identification. Measurements were, however, made of one 
hundred trypanosomes, fifty from hump-backed cattle, and the same 
number from straight-backed cattle (see Table IV). The average 
length was found to be 22 64 /x, and the range from 18// to 29 yu. 
These measurements, and the curve shown in Chart II, are 
consistent with the identification T. vivax. 


Table IV.—Measurements in length of T. vivax from cattle. 


Host 

Number 

measured 

Distribution according to length in microns 

Average 

18 

O 

21 

22 

23 

24 


26 ; 27 

28 

29 

Cattle, hump-backed, No. 451 ... 

25 


I 2 

+ 

4 

8 

5 

1 

! 

— 

— 

22*40 

Cattle, hump-backed, No. 630 ... 

*5 


I 1 — 

I 

4 

9 

6 

1 

2 1 

— 

— 

23*32 

Cattle, straight-backed, No. 589 

25 

1 

— 1 2 

2 

5 

5 

3 

4 

1 1 1 

— 

1 

23*16 

Cattle, straight-backed, No. 734 

25 

— 

— I 

5 

5 

7 

4 

3 

~~ “ 

— 

— 

22*68 


100 

1 

' 

2 5 

1 

12 

18 

29 

1- 

9 

3 2 

— 

1 

22*64 






47i 



Chart II.—The distribution, by percentages, in length of T. vivax 

from cattle. 





















472 


T. congolense 

Small monomorphic trypanosomes with a scanty undulating 
membrane, and without a free portion to the flagellum, were present 
in forty-two (8 4 per cent.) of the animals examined, namely, in 
twenty-eight hump-backed cattle, six straight-backed cattle, two 
sheep, five pigs, and one goat. The infections were never very 
heavy, and in several of the hosts only one or two individuals were 
found in a large blood-film. 

Both a guinea-pig and a white rat were on two occasions 
inoculated with blood containing these small trypanosomes, but 
with negative results. It was unfortunately impossible to experiment 
with larger animals, such as goats, sheep, and cattle, and no dogs 
were available. No detailed measurements at known periods of the 
infection could therefore be made. Fifty individuals, taken as they 
came, were, however, measured in blood-films from hump-backed 
cattle and pigs naturally infected with these parasites (see 
Table V). The fifty from cattle averaged 12*941* in length; the 
longest measuring 17/* and the shortest 10 a*. The average breadth 
was 2*29 a*, the broadest being 3*5/1 and the narrowest 1*25 a*. In 
pigs the average length of the fifty trypanosomes measured was 
12 24A*, the longest being 16/1 and the shortest ioa*. In breadth 
they averaged 199/*, and ranged from 3*25/* to 125/*. The 
distribution according to length and breadth of the hundred 
individuals is shown in Charts III and IV. 

The identity of these trypanosomes is open to doubt. Similar 
specimens from horses which were taken to England from Nigeria in 
1911 were identified by Bruce as T. pecorum , a species which is 
apparently indistinguishable from T. nanunt , and which, according 
to Blacklock and Yorke (1913), is probably identical with 
T. congolense. The trypanosome found in the Gold Coast is 
morphologically similar to that found in Nigeria, although in both 
colonies the parasite was found to vary from the type to a consider¬ 
able extent. In experiments conducted at Eket this variation was 
found to occur in the same strain when transmitted to different 
hosts. There seems to be some reason to suppose that T . fecorum> 
T . nanunt , and T. congolense are all strains of the same species of 
trypanosome; in which case the name T. congolense is that properly 
applicable to it. 



473 


With regard to T. dimorphon , such uncertainty exists as to the 
nature of the original strain that it is doubtful if the name should 
now be employed. The original figures given by Dutton and Todd 
(1903) of their Gambian Horse Trypanosome show two types of 
parasite that would certainly to-day be identified in Nigeria or the 
Gold Coast as T. congolense (T. nanuni ) and T. pecaudi (T . brucei 
of Uganda). An infected horse sent home some years later by 

Table V.—The measurements in length and breadth of T. congolense ( T . nanum ). 


Length in microns Distribution according to length in microns 

Host Number _ _ 



measured 

Max. 

Average 

1 Min. 

i 

8 

9 

10 

11 

12 

>3 

14 

*5 

16 

17 

18 

Pigs. 

... 50 

16 

12-24 

| 10 

— 

— 

2 

11 

*3 

9 

7 

6 

2 

— 

— 

Cattle, humped 

50 

*7 

I2 ’94 

IO 

— 

— 

9 

2 

10 

9 

11 

3 

3 

3 

— 


100 

l 7 

! 2-59 

10 

— 

— 

11 

1 13 

2 3 

18 

18 

9 

5 

3 

— 


Host 

, Number 

Breadth in Microns 

Distribution 

according to breadth in microns 



measured 

Max. 

1 Average 

Min. 1 

1 * ^ 

«-5 

i *75 

2 

2-25 

1.5 

2-75 

3 

3* 2 5 

3*5 

Pigs. 

... 50 

3*25 

i -99 

1-25 

— 

4 

4 

" 1 

*4 

8 

3 

4 

— 

1 

— 

Cattle, humped 

... 50 

3-5 

2*29 

1*25 

— 

1 

3 

3 

! 5 

5 

12 

2 

6 

2 1 

1 


100 

3-5 

| 

2-24 

1 *25 


5 

7 ' 

15 

2 9 

*3 

*5 

6 

6 

_j 

3 

1 


Dr. Todd as probably harbouring T. dimorphon proved on investi¬ 
gation by Yorke and Blacklock to have a double infection with 
T. congolense and T. vivax. It is probable, therefore, that the 
animals originally examined by Dutton and Todd were infected 
with all the three types of trypanosome common in West Africa, 
namely, those referred to here as T. vivax , T. congolense , and 
T. pecaudi. In the sense in which the name is used by Laveran and 
Mesnil, T. dimorphon denotes a polymorphic species of trypano- 





474 



Chart III.—The distribution, by percentages, in length of T. congolense 

(T. nanutri). 









475 



Chart IV.—The distribution, by percentages, in breadth of T. congoUnse 

(T. nanum ). 









476 


some, highly pathogenic to the smaller laboratory animals, all the 
individuals of which are without a free flagellum, and which shows 
forms measuring from 11 fi up to 30 fi in length. Such a trypanosome 
I have not yet seen in West Africa. Mesnil (1915) considers that the 
small monomorphic strains of trypanosome isolated by me at Eket 
from G . tachinoides were T. dtmorphon , apparently because a single 
individual was found to attain the length of 21//. Two strains were 
isolated by me, and in the second a single trypanosome was 
measured that reached this length. This strain I noted in my 
description was unusually large, but I did not separate it off from 
the other and more typical one because I observed considerable 
variations in the size of the latter parasite when inoculated into 
different animals. The trypanosomes of the first and more typical 
strain measured from 9/1 to 18/*, and averaged 13*9/* in length; 
and I think it could not but have been T. congoletise. If the second 
strain were not a variety of the same parasite, as I believe it to have 
been, it must, I think, have been a new species. Its length, average 
15’ 15A 1 , maximum 21/1, minimum n /*, does not approach that of 
T. dimorphon. 

The ideal relationship between a parasite and its host would 
appear to be the benign infection, since a strain that kills its host 
commits racial suicide. In the case of these trypanosomes an 
approximation to the ideal relationship exists in game, and in 
certain domestic animals. It might be expected, however, that a 
parasite specialised so as to produce a benign infection in a certain 
type of host might show wide variations in pathogenicity when 
introduced into other animals. The smaller laboratory animals, 
such as rats, guinea-pigs, and rabbits, in which the pathogenicity 
of trypanosoifies is generally studied, are species that tsetse flies 
normally would seldom have an opportunity of infecting, and